There are few scientists of whom it can be said that their mistakes are more int

eresting
than their colleagues' successes, but Albert Einstein was one. Few "blunders" ha
ve had a
longer and more eventful life than the cosmological constant, sometimes describe
d as the
most famous fudge factor in the history of science, that Einstein added to his t
heory of
general relativity in 1917. Its role was to provide a repulsive force in order t
o keep the
universe from theoretically collapsing under its own weight. Einstein abandoned
the
cosmological constant when the universe turned out to be expanding, but in succe
eding
years, the cosmological constant, like Rasputin, has stubbornly refused to die,
dragging
itself to the fore, whispering of deep enigmas and mysterious new forces in natu
re,
whenever cosmologists have run into trouble reconciling their observations of th
e
universe with their theories.
This year the cosmological constant has been propelled back into the news as an
explanation for the widely reported discovery, based on observations of distant
exploding
stars, that some kind of "funny energy" is apparently accelerating the expansion
of the
universe. "If the cosmological constant was good enough for Einstein," the cosmo
logist
Michael Turner of the University of Chicago remarked at a meeting in April, "it
should
be good enough for us."
Einstein has been dead for 43 years. How did he and his 80-year-old fudge factor
come to
be at the center of a revolution in modern cosmology?
The story begins in Vienna with a mystical concept that Einstein called Mach's p
rinciple.
Vienna was the intellectual redoubt of Ernst Mach (1838-1916), a physicist and
philosopher who bestrode European science like a Colossus. The scale by which
supersonic speeds are measured is named for him. His biggest legacy was philosop
hical;
he maintained that all knowledge came from the senses, and campaigned relentless
ly
against the introduction of what he considered metaphysical concepts in science,
atoms
for example.
1Another was the notion of absolute space, which formed the framework of Newton'
s
universe. Mach argued that we do not see "space," only the players in it. All ou
r
knowledge of motion, he pointed out, was only relative to the "fixed stars." In
his books
and papers, he wondered if inertia, the tendency of an object to remain at rest
or in
motion until acted upon by an outside force, was similarly relative and derived
somehow
from an interaction with everything else in the universe.
"What would become of the law of inertia if the whole of the heavens began to mo
ve and
stars swarmed in confusion?" he wrote in 1911. "Only in the case of a shattering
of the

universe do we learn that all bodies, each with its share, are of importance in
the law of
inertia."
Mach never ventured a guess as to how this mysterious interaction would work, bu
t
Einstein, who admired Mach's incorrigible skepticism, was enamored of what he
sometimes called Mach's principle and sometimes called the relativity of inertia
. He
hoped to incorporate the concept in his new theory of general relativity, which
he
completed in 1915. That theory describes how matter and energy distort or "curve
" the
geometry of space and time, producing the phenomenon called gravity.
In the language of general relativity, Mach's principle required that the spacetime
curvature should be determined solely by other matter or energy in the universe,
and not
any initial conditions or outside influences -- what physicists call boundary co
nditions.
Among other things, Einstein took this to mean that it should be impossible to s
olve his
equations for the case of a solitary object -- an atom or a star alone in the un
iverse -since there would be nothing to compare it to or interact with.
So Einstein was surprised a few months after announcing his new theory, when Kar
l
Schwarzschild, a German astrophysicist serving at the front in World War I, sent
him just
such a solution, which described the gravitational field around a solitary star.
"I would
not have believed that the strict treatment of the point mass problem was so sim
ple,"
Einstein said.
Perhaps spurred in part by Schwarzschild's results, Einstein turned his energies
in the fall
of 1916 to inventing a universe with boundaries that would prevent a star from e
scaping
its neighbors and drifting away into infinite un-Machian loneliness. He worked o
ut his
ideas in a correspondence with a Dutch astronomer, Willem de Sitter, which are t
o be
published this summer by the Princeton University Press in Volume 8 of "The Coll
ected
Papers of Albert Einstein." Like most of his colleagues at the time, Einstein co
nsidered
2the universe to consist of a cloud of stars, namely the Milky Way, surrounded b
y vast
space. One of his ideas envisioned "distant masses" ringing the outskirts of the
Milky
Way like a fence. These masses would somehow curl up space and close it off.
His sparring partner de Sitter scoffed at that, arguing these "supernatural" mas
ses would
not be part of the visible universe. As such, they were no more palatable than N
ewton's
old idea of absolute space, which was equally invisible and arbitrary.
In desperation and laid up with gall bladder trouble in February of 1917, Einste
in hit on
the idea of a universe without boundaries, in which space had been bent around t
o meet
itself, like the surface of a sphere, by the matter within. "I have committed an

other
suggestion with respect to gravitation which exposes me to the danger of being c
onfined
to the nut house," he confided to a friend.
This got rid of the need for boundaries -- the surface of a sphere has no bounda
ry. Such a
bubble universe would be defined solely by its matter and energy content, as Mac
hian
principles dictated. But there was a new problem; this universe was unstable, th
e bubble
had to be either expanding or contracting. The Milky Way appeared to be neither
expanding nor contracting; its stars did not seem to be going anywhere in partic
ular.
Here was where the cosmological constant came in. Einstein made a little mathema
tical
fix to his equations, adding "a cosmological term" that stabilized them and the
universe.
Physically, this new term, denoted by the Greek letter lambda, represented some
kind of
long range repulsive force, presumably that kept the cosmos from collapsing unde
r its
own weight.
Admittedly, Einstein acknowledged in his paper, the cosmological constant was "n
ot
justified by our actual knowledge of gravitation," but it did not contradict rel
ativity,
either. The happy result was a static universe of the type nearly everybody beli
eved they
lived in and in which geometry was strictly determined by matter. "This is the c
ore of the
requirement of the relativity of inertia," Einstein explained to de Sitter. "To
me, as long
as this requirement had not been fulfilled, the goal of general relativity was n
ot yet
completely achieved. This only came about with the lambda term."
The joke, of course, is that Einstein did not need a static universe to have a M
achian one.
Michel Janssen, a Boston University physicist and Einstein scholar, pointed out,
"Einstein needed the constant not because of his philosophical predilections but
because
of his prejudice that the universe is static."
3Moreover, in seeking to save the universe for Mach, Einstein had destroyed Mach
's
principle. "The cosmological term is radically anti-Machian, in the sense that i
t ascribes
intrinsic properties (energy and pressure-density) to pure space, in the absence
of matter,"
said Frank Wilczek, a theorist at the Institute for Advanced Study in Princeton.
In any event, Einstein's new universe soon fell apart. In another 10 years the a
stronomer
Edwin Hubble in California was showing that mysterious spiral nebulae were galax
ies far
far away and getting farther -- in short that the universe might be expanding.
De Sitter further confounded Einstein by coming up with his own solution to Eins
tein's
equations that described a universe that had no matter in it at all.
"It would be unsatisfactory, in my opinion," Einstein grumbled, "if a world with
out
matter were possible."
De Sitter's empty universe was also supposed to be static, but that too proved t

o be an
illusion. Calculations showed that when test particles were inserted into it, th
ey flew
away from each other. That was the last straw for Einstein. "If there is no quas
i-static
world," he said in 1922, "then away with the cosmological term."
In 1931, after a trip to the Mount Wilson observatory in Pasadena, Calif., to me
et Hubble,
Einstein turned his back on the cosmological constant for good, calling it "theo
retically
unsatisfactory anyway."
He never mentioned it again.
In the meantime, the equations for an expanding universe had been independently
discovered by Aleksandr Friedmann, a young Russian theorist, and by the Abbe Geo
rges
Lemaitre, a Belgian cleric and physicist. A year after his visit with Hubble, Ei
nstein
threw his weight, along with de Sitter, behind an expanding universe without a
cosmological constant.
But the cosmological constant lived on in the imagination of Lemaitre, who found
that by
judicious application of lambda he could construct universes that started out ex
panding
slowly and then sped up, universes that started out fast and then slowed down, o
r one that
even began expanding, paused, and then resumed again.
This last model beckoned briefly to some astronomers in the early 1950's, when
measurements of the cosmic expansion embarrassingly suggested that the universe
was
4only two billion years old -- younger Earth. A group of astronomers visited Ein
stein in
Princeton and suggested that resuscitating the cosmological constant could resol
ve the
age discrepancy. Einstein turned them down, saying that the introduction of the
cosmological constant had been the biggest blunder of his life. George Gamow, on
e of
the astronomers, reported the remark in his autobiography, "My World Line," and
it
became part of the Einstein legend.
Einstein died three years later. In the years after his death, quantum mechanics
, the
strange set of rules that describe nature on the subatomic level (and Einstein's
bete noire)
transformed the cosmological constant and showed just how prescient Einstein had
been
in inventing it. The famous (and mystical in its own right) uncertainty principl
e decreed
that there is no such thing as nothing, and even empty space can be thought of a
s foaming
with energy.
The effects of this vacuum energy on atoms had been detected in the laboratory,
as early
as 1948, but no one thought to investigate its influence on the universe as a wh
ole until
1967, when a new crisis, an apparent proliferation of too-many quasars when the
universe
was about one-third its present size, led to renewed muttering about the cosmolo
gical
constant. Jakob Zeldovich, a legendary Russian theorist who was a genius at marr
ying

microphysics to the universe, realized that this quantum vacuum energy would ent
er into
Einstein's equations exactly the same as the old cosmological constant.
The problem was that a naive straightforward calculation of these quantum fluctu
ations
suggested that the vacuum energy in the universe should be about 118 orders of
magnitude (10 followed by 117 zeros) denser than the matter. In which case the
cosmological constant would either have crumpled the universe into a black hole
in the
first instant of its existence or immediately blown the cosmos so far apart that
not even
atoms would ever have formed. The fact that the universe had been sedately and h
appily
expanding for 10 billion years or so, however, meant that any cosmological const
ant, if it
existed at all, was modest.
Even making the most optimistic assumptions, Dr. Zeldovich still could not make
the
predicted cosmological constant to come out to be less than a billion times the
observed
limit.
Ever since then, many particle theorists have simply assumed that for some as-ye
tunknown reason the cosmological constant is zero. In the era of superstrings and
ambitious theories of everything tracing history back to the first micro-micro s
econd of
unrecorded time, the cosmological constant has been a trapdoor in the basement o
f
5physics, suggesting that at some fundamental level something is being missed ab
out the
world. In an article in Reviews of Modern Physics in 1989, Steven Weinberg of th
e
University of Texas referred to the cosmological constant as "a veritable crisis
," whose
solution would have a wide impact on physics and astronomy.
Things got even more interesting in the 1970's with the advent of the current cr
op of
particle physics theories, which feature a shadowy entity known as the Higgs fie
ld, which
permeates space and gives elementary particles their properties. Physicists pres
ume that
the energy density of the Higgs field today is zero, but in the past, when the u
niverse was
hotter, the Higgs energy could have been enormous and dominated the dynamics of
the
universe. In fact, speculation that such an episode occurred a fraction of a sec
ond after
the Big Bang, inflating the wrinkles out of the primeval chaos -- what Dr. Turne
r calls
vacuum energy put to a good use -- has dominated cosmology in the last 15 years.
"We want to explain why the effective cosmological constant is small now, not wh
y it
was always small," Dr. Weinberg wrote in his review. In their efforts to provide
an
explanation, theorists have been driven recently to talk about multiple universe
s
connected by space-time tunnels called wormholes, among other things.
The flavor of the crisis was best expressed, some years ago at an astrophysics c
onference
by Dr. Wilczek. Summing up the discussions at the end of the meeting, he came at

last to
the cosmological constant. "Whereof one cannot speak, thereof one must be silent
," he
said, quoting from Ludwig Wittgenstein's "Tractatus Logico-Philosophicus."
Now it seems that the astronomers have broken that silence.
Copyright 2002 The New York Times Company
6Mysteries of the Universe
Q U A N T U M P H Y S I C S
Quantum Theory Tugged, and All of Physics Unraveled
By DENNIS OVERBYE
They tried to talk Max Planck out of becoming a physicist, on the grounds that h
ere was
nothing left to discover. The young Planck didn't mind. A conservative youth fro
m the
south of Germany, a descendant of church rectors and professors, he was happy to
add to
the perfection of what was already known.
Instead, he destroyed it, by discovering what was in effect a loose thread that
when
tugged would eventually unravel the entire fabric of what had passed for reality
.
As a new professor at the University of Berlin, Planck embarked in the fall of 1
900 on a
mundane sounding calculation of the spectral characteristics of the glow from a
heated
object. Physicists had good reason to think the answer would elucidate the relat
ionship
between light and matter as well as give German industry a leg up in the electri
c light
business. But the calculation had been plagued with difficulties.
Planck succeeded in finding the right formula, but at a cost, as he reported to
the German
Physical Society on Dec. 14. In what he called "an act of desperation," he had t
o assume
that atoms could only emit energy in discrete amounts that he later called quant
a (from
the Latin quantus for "how much" ) rather than in the continuous waves prescribe
d by
electromagnetic theory. Nature seemed to be acting like a fussy bank teller who
would
not make change, and would not accept it either.
That was the first shot in a revolution. Within a quarter of a century, the comm
on sense
laws of science had been overthrown. In their place was a bizarre set of rules k
nown as
quantum mechanics, in which causes were not guaranteed to be linked to effects;
a
subatomic particle like an electron could be in two places at once, everywhere o
r
nowhere until someone measured it; and light could be a wave or a particle.
and the unknown, who as been accorded some of the ultimate accolades in pop cult
ure -appearing as Einstein's poker buddy on "Star Trek: The Next Generation," and as
a guest
star on "The Simpsons."
While a graduate student, in 1963, he learned he had amyotrophic lateral scleros
is and
was given a few years to live. He has moved about in a wheelchair for more than
25 years

and now speaks only through a voice synthesizer. Dr. Hawking, for whom the word
"puckish" seems to have been invented, has often said his disability is an advan
tage
because it frees him to sit and think. Next month his colleagues will celebrate
his 60th
birthday with a weeklong all-star symposium in Cambridge.
In the new book's introduction, Dr. Hawking admits that "A Brief History of Time
" was
"not easy going" and laments that some readers got stuck and did not finish it.
He has
tried, he says, to make this one easier. Slightly longer than the earlier book,
"Nutshell," at
216 pages, is embellished with colorful illustrations that give it a coffee-tabl
e-book look.
So far the critics are in qualified agreement; one, Bryan Appleyard in the The N
ew
he Bivalvia, the second largest class within the
Solnhofen Limestone of Eichsta tt, Germany, and was
described by Cosimo Collini (17271806) in 1784.
Collini concluded that it was a possible sea creature
of unknown affinity, although he did note bat-like
features. In 1801, the great French anatomist Georges
Cuvier (17691832) recognized that the creature
was a reptile and that its elongated digits must have
supported flight membranes. Cuvier was thus the first
to recognize pterosaurs as flying reptiles and, in 1809,
he coined the name Ptero-Dactyle. This later became
the generic name Pterodactylus (Figures 1 and 4).
In the decades that followed, a succession of further
pterosaurs from the Solnhofen Limestone was announced, many in a spectacular state of preservation
and some with their wing membranes intact. The
first recognized British pterosaur, a specimen of the
deep-skulled Dimorphodon, was discovered by Mary
Anning (17991847) in 1827 in Lower Jurassic rocks
of Lyme Regis, Dorset. We now know that Gideon
Mantell (17901852), best known for the discovery
of Iguanodon, found pterosaur remains before this
in the Early Cretaceous Wealden strata of Sussex,
but had thought that these were from birds. North
America yielded its first pterosaur to the prolific
palaeontologist O. C. Marsh (18311899) in 1871
and, by 1876, Marsh had recognized it as a new,
distinctive genus he named Pteranodon (meaning
winged and toothless). With an estimated wingspan
of 6 m, Pteranodon was huge compared to most
earlier discoveries.
While these discoveries and others were being
made, varied opinions on the nature and life style of
pterosaurs were appearing, and they were variously
depicted as swimming creatures, as bats, marsupials,
or as kin of birds. By the early 1900s, it was generally
agreed that pterosaurs were bat-like flying reptiles and,
in 1901, Harry Seeley (18391909) published Dragons
of the Air, the first book devoted to pterosaurs.
South American Cretaceous pterosaurs have proved
to be among the most important in the world, but not
until 1971 was the first pterosaur from the now famous
Santana Formation of Brazil discovered. Since then a
significant number of new kinds from around the

world (around 70 genera are presently recognized)

have revealed previously unimagined morphologies
and maximum sizes. Until 1971, Pteranodon sternbergi
Figure 1 Life restoration of the Late Jurassic pterodactyloid, Pterodactylus , f
rom the German Solnhofen Limestone in a quadrupedal
stance. Note the presence of body hair and the soft tissue head crest. Reproduce
d with permission from Dino Frey. Buffetaut E
and Mazin J M (2003) Evolution and Palaeobiology of Pterosaurs. Geological Socie
ty Special Publication 217 . London: The Geological Society
of London.510 FOSSIL VERTEBRATES/Flying Reptiles
(wingspan, 9 m) was the largest known flying animal,
but the discovery in Texas of Quetzalcoatlus revealed
that the biggest pterosaurs achieved wingspans of
11 m. Related pterosaurs of similar or larger size were
discovered in the 1990s in Spain and eastern Europe.
The Pterosaur Skeleton
The pterosaur skeleton was highly modified for flight,
and the most obvious features are the huge size of the
skull compared with the body and the extreme
elongation of one of the fingers. Like birds, most
pterosaurs had hollow bones with foramina (small
openings), indicating that they contained air sacs
connected to the lungs. Pterosaur bones were supported internally by struts, and the bone walls themselves, usually no thicker than 2 mm, are composed
of multiple overlapping layers and thus combine
lightness with strength.
Pterosaur skull morphology is varied, although the
majority had long, slim, shallow jaws and all had
large orbits (eye-sockets). In basal pterosaurs, the
external nostril was separate from an opening in
front of the orbit called the antorbital fenestra. In
pterodactyloids, these two openings merged into a
single one called the nasoantorbital fenestra. Pterosaur teeth were extremely variable. Widely spaced
pointed teeth, were widespread and from ancestors
with teeth like these evolved species with fang-like
teeth at the jaw tips and the unique Istiodactylus
with its short petal-shaped teeth. The Late Triassic
Eudimorphodon and Austriadactylus possessed
multicusped teeth while elongate, slender teeth
numbering in the hundreds evolved in the ctenochasmatoids. Toothlessness evolved several times. Some
pterosaurs skulls sport bony crests at the jaw tips,
along the midline or at the back of the skull.
Figure 2 Wing skeleton of an ornithocheiroid pterodactyloid.
Unlike birds and bats, the main wing spar in pterosaurs was formed by a hypertrophied digit (Figure 2).
This wing finger is generally considered to be the
fourth because the digital formulae of the pterosaur
hand best matches that of digits one to four in the
hands of other reptiles. However, a rod-shaped bone
projecting from the pterosaur wrist, called the pteroid
bone, has at times been argued to represent the first
hand digit. This is a minority view today but, if it is
correct, then pterosaurs have five hand digits and the
wing finger is the fifth. Although most pterosaur
fossils show the pteroid pointing towards the shoulder, some workers suggest that it pointed forwards
parallel to the neck. Regardless, the pteroid was

probably mobile and used to control the attitude of

the propatagium (Figure 3).
The pterosaur pectoral girdle includes a (normally
fused) scapula and coracoid that meet at an acute
angle and, as expected for flying animals, the socket
for the humerus faces sideways and slightly upwards.
The coracoids attach to an enlarged keeled sternum
that anchored most of the major flight muscles. The
bones of the pterosaur pelvis were short, usually fused
together, and with a closed hip socket. Pterosaurs
have a pair of unique rod- or plate-like bones called
the prepubes projecting forwards from the bottom
of the pelvis. Like the gastralia (belly ribs) that all
pterosaurs possessed, they may have helped support
the gut or keep the abdomen rigid.
The vertebral column in pterosaurs can clearly be
differentiated into cervical, dorsal, sacral, and caudal
portions. The number of vertebrae is variable and
pterosaurs have 79 cervical, 1116 dorsal, 310
sacral, and 1140 caudal vertebrae. The many caudal
vertebrae of basal pterosaurs are encased in long bony
processes that make the tail stiff and rod-like. In
derived Cretaceous pterosaurs, most of the dorsalFOSSIL VERTEBRATES/Flying Repti
les 511
vertebrae are fused together forming a structure
called the notarium.
Compared with the other wing segments, the pterosaur humerus is short, although generally with massive crests for muscle attachment. The ulna is always
larger than the radius and both are attached distally
to block-shaped carpal bones. Projecting from one
of these is the unique pteroid. Pterosaurs had four
metacarpals, the first three of which were slim and,
with the exception of Nyctosaurus from Late Cretaceous North America, attached to short, clawed
fingers. Why Nyctosaurus lacked clawed fingers is
unknown, but in all other pterosaurs these digits
may have served important functions. Trackways
show that they were used in walking, and it is also
possible that they were employed in grooming or
climbing. The fourth metacarpal was robust and
tipped with a twisted, roller-like distal end to which
was attached the massive wing finger. This consists of
four long straight bones, excepting a few genera
where there were only three. Because of the twisted
end of the fourth metacarpal, the wing finger would
have lain parallel to the bodys long axis when the
wing was folded up.
The pterosaur hind limb is lightly built and the
head of the femur is only slightly offset from the
long axis of the shaft. Pterosaur hind limbs seem to
have been quite flexible, but mostly sprawled to the
sides. During flight, the hind limb was probably held
in a bat-like orientation and could have been used to
control the shape of the wing membranes. Basal
pterosaurs are five-toed, with a prominent curving
Figure 3 Schematic representation of the flight membranes in
a generalized pterodactyloid pterosaur.
fifth digit that is hooked towards the tail. In pterodactyloids, the fifth digit is either absent or present as

a tiny stub.
Because some articulated fossils indicate that the
foot could assume a 90  angle relative to the tibia
(and there is little evidence for much motion at the
metatarsophalangeal joints), pterosaurs have generally been regarded as plantigrade (placing the whole
length of the foot on the ground when walking). In
1983, Kevin Padian argued that this was not the case
for Dimorphodon and that it may instead have been
digitigrade (walking only on the toes). This was later
inferred for all pterosaurs. An articulated Dimorphodon foot shows, however, that only limited motion
was possible at the metatarsophalangeal joint, thus
supporting a plantigrade posture. This is in agreement
with probable pterosaur tracks preserved as trace
fossils.
Soft Tissue, Integument, and
Pterosaur Life Appearance
Many aspects of pterosaur life appearance remain
unknown or controversial, although a number of exceptional fossils have provided some surprising
details. Pterosaur body hair was reported as early as
1831 and described for various Jurassic pterosaurs
between the 1920s and 1970s and today it is clear
that pterosaurs had bristle-like hairs covering their
necks and bodies (Figure 4). The active flapping flight
and body hair of pterosaurs suggest that they had an
elevated metabolism.
Other exceptional fossils show that some pterosaurs possessed a throat pouch, webbing between
the toes, and scales on the soles of the feet. Soft
Figure 4 An exceptionally well preserved skeleton of the Late
Jurassic pterodactyloid Pterodactylus from the German Solnhofen
Limestone. This specimen preserves parts of the flight mem
branes, a throat pouch, and hairs on the neck and back.512 FOSSIL VERTEBRATES/Fl
ying Reptiles
Figure 5 Variation in skull crest morphology in pterodactyloids. Soft tissue cre
sts are now known for a wide diversity of
pterodactyloids. Reproduced with permission from Dino Frey and Marie Celine Buch
y. Buffetaut E and Mazin J M (2003) Evolution
and Palaeobiology of Pterosaurs. Geological Society Special Publication 217 . Lo
ndon: The Geological Society of London.
tissue skull crests connected to the underlying bony
crests have proved to be widespread and appear
to have doubled the size of the bony crests (Figures 5
and 6). An unexpected discovery is a soft tissue crest
in Pterodactylus, a genus that lacks a bony crest
(Figure 1). The presence of a distinctive bone texture
on the pterosaur snout, jaw, and palate indicates that
pterosaurs were beaked.
Pterosaur wing membranes are known from wellpreserved specimens from the Solnhofen Limestone
and the Early Cretaceous Brazilian Crato and Santana formations. A membrane called the propatagium
extended from the shoulder to the pteroid and perhaps distally to encompass the first three fingers. The
main flight membrane, the brachiopatagium (also
called the cheiropatagium), extended from the tip of
the wing finger to the hind limb, extending
as far distally as the knee, shin, or ankle. Another

membrane, the uropatagium, was present between

the hind limbs (Figure 3). The wing membrane
appears to have been complex, with a thin epidermis,
a layer of vascular tissues, a layer of stiffening fibres
called aktinofibrils, a thin sheet of muscle, and a
Figure 6 Skull of the tapejarid pterosaur Topejara navigaus from
the Early Cretaceous Crato Formation of Brazil with bony and soft
tissue skull crest. Buffetaut E and Mazin J M (2003) Evolution and
Palaeobiology of Pterosaurs. Geological Society Special Publication 217 .
London: The Geological Society of London.FOSSIL VERTEBRATES/Flying Reptiles 513
blood capillary network. Rhamphorhynchus and
probably other long-tailed pterosaurs possessed a
vertical diamond-shaped membrane at the tail tip.
With skin membranes connecting the wings, body,
and legs, pterosaurs may have been superficially batlike but, because bats are mostly dark-coloured nocturnal animals, it is doubtful that the similarities were
strong. Pterosaurs mostly seem to have been ecological
analogues of sea- and water-birds, and it might be
that they were patterned in whites, blacks, and greys,
although bright colours presumably decorated their
crests.
pterosaur hind limbs are only superficially similar to
those of dinosaurs, and that re-analysis favoured a
position for pterosaurs outside of the crocodilian
dinosaur group. Rather more heterodox recent ideas
include the suggestion that pterosaurs are the closest
relatives of birds and that pterosaurs are part of the
Dinosauria.
Several different models have been proposed for
the origin of pterosaurs, but the presence in basal pterosaurs of climbing features and of various details in the
hind limb and pelvis indicative of a leaping ability suggest that pterosaurs first evolved as tree-climbing
leapers.
The Affinities and Origin of Pterosaurs
Historically, pterosaurs have been allied with Mesozoic marine reptiles, bats, marsupials, and birds
(see Fossil Vertebrates: Dinosaurs; Birds). However,
major improvements in the understanding of vertebrate evolution allowed the palaeontologists of the
nineteenth and twentieth centuries to realize that
pterosaurs were related at least vaguely to dinosaurs
(see ) and their allies.
Although it is clear that pterosaurs are part of the
major reptile assemblage known as the Diapsida,
their affinities within this group are controversial.
The presence of an antorbital fenestra has conventionally meant that pterosaurs have been regarded
as archosaurs, the so-called ruling reptile group
that incorporates crocodilians, dinosaurs, and kin.
Among archosaurs, pterosaurs share a simple hingelike ankle joint with dinosaurs and consequently
have been regarded as close relatives of dinosaurs in
most studies. This view was developed at a time when
some workers thought that pterosaurs originated
from terrestrial bipedal ancestors and that pterosaurs
themselves were bipedal and digitigrade. A small bipedal, long-legged archosaur from Late Triassic
Scotland, Scleromochlus, was argued to be a ptero-

saur ancestor, but recent studies refute this idea. The

idea that pterosaurs might be close relatives of dinosaurs can certainly be regarded as the mainstream
view in vertebrate palaeontology today. However,
several recent studies have questioned the evidence
for this proposed affinity.
An alternative hypothesis argues that pterosaurs
belong instead to a group of archosaur-like diapsids,
the Prolacertiformes. Most prolacertiforms were
superficially lizard-like, but Sharovipteryx from Late
Triassic Kyrgyzstan appears to be intermediate between conventional prolacertiforms and pterosaurs.
It has pterosaur-like hind limbs and vertebrae and
membranes between its hind limbs and tail.
Some other models for pterosaur ancestry have been
proposed. In 1996, S. Christopher Bennett argued that
Pterosaur Diversity and Phylogeny
It was recognized in 1901 that pterosaurs could be
divided into two groups: the toothed, mostly longtailed Rhamphorhynchoidea, and the short-tailed
Pterodactyloidea (including both toothed and toothless kinds). Today, it is clear that rhamphorhynchoids
include the ancestors of pterodactyloids and, consequently, Rhamphorhynchoidea is a grade and not a
clade. Pterosaurs previously referred to as rhamphorhynchoids are nowadays termed basal pterosaurs or
non-pterodactyloids. Although basal pterosaurs were
diverse, it is notable that they were small compared
with the majority of Cretaceous pterodactyloids.
The evolutionary relationships of pterosaurs are
relatively understudied and only recently has pterosaur
phylogeny been analysed. Although some areas of consensus have emerged, authors disagree on the details.
We follow the phylogeny proposed by David Unwin
of the Museum fu r Naturkunde in Berlin (Figure 7).
Perhaps the most basal pterosaur is Preondactylus
from the Late Triassic of Italy. This form has a shorter
coracoid and humerus and longer legs than other
pterosaurs. Dimorphodontids, which include Dimorphodon from Early (and perhaps Middle) Jurassic
Figure 7 Cladogram depicting the relationships of all the major
pterosaur groups. Reproduced from David Unwin.514 FOSSIL VERTEBRATES/Flying Rept
iles
England and Mexico, are basal pterosaurs with deep
skulls superficially like those of puffins, whilst anurognathids were unusual in having short, broad snouts
and abbreviated tails. A surprising recent discovery
is the persistence of anurognathids into the Early
Cretaceous. Two basal pterosaurs, Eudimorphodon
from Late Triassic Italy and Greenland and Campylognathoides from Early Jurassic Germany and India,
are united in the Campylognathoididae based on a
distinctive lower jaw in which two pairs of large
conical teeth are followed by multiple smaller ones.
Rhamphorhynchids were successful Jurassic pterosaurs known from Eurasia, North America, and
Africa. Rhamphorhynchus from Late Jurassic Europe
exhibits a laterally compressed, ventrally directed
lower jaw tip and an array of forward-pointing
teeth. It probably used these to grab fish and other

small prey from the water. Another rhamphorhynchid

lineage, the scaphognathines, had deeper skulls with
teeth perpendicular to the jaw margins.
The Pterodactyloids
Pterodactyloids, the advanced short-tailed pterosaurs,
originated from a rhamphorhynchid-like ancestor
during the Middle Jurassic. The pterodactyloid radiation consisted of four major groups: the robust-jawed
ornithocheiroids, the slim-jawed ctenochasmatoids,
the low-crested dsungaripteroids, and the long-necked,
crested azhdarchoids. A fifth group, the lonchodectids
from Early Cretaceous England, are of uncertain affinity. Lonchodectids were small (wingspan, 12 m) with
long, dorsoventrally flattened jaws with small teeth,
each of which was supported by a low bony collar at
its base.
Ornithocheiroids were large predatory pterosaurs
(wingspan, 29 m) with robust beaks, often housing
recurved, fang-like teeth at their tips (Figure 8). Their
jaws frequently possessed keel-like dorsal and ventral
crests, and some forms also possessed crests on the
back of the skull. The toothless pteranodontids and
nyctosaurids appear to be members of this group. The
earliest known ornithocheiroids appear at the start of
the Cretaceous, while nyctosaurids survived to the
very end of the Cretaceous.
Ctenochasmatoids had needle-like meshes of teeth
set in long, thin jaws. In Pterodaustro from late
Early Cretaceous Argentina, the upturned lower jaw
contains approximately 1000 bristle-like teeth. These
were surely used for filtering small organisms from
the water. Unlike ornithocheiroids, ctenochasmatoids
had elongate cervical vertebrae and were generally
small (wingspan, 50 cm to 2 m), although Cearadactylus from Early Cretaceous Brazil was a giant with
a wingspan of 5.5 m.
Figure 8 Jaw tips of the Cretaceous ornithocheiroid pterosaur
Coloborhynchus . Note the massive fang like anterior teeth and the
low keel like crests on both jaws. This specimen is from the
Santana Formation of Brazil and would have belonged to an
animal with a skull of approximately 1 m in length.
Members of the Dsungaripteroidea are known from
the Late Jurassic and Early Cretaceous of Eurasia,
Africa, and South America. Like some ctenochasmatoids, dsungaripteroids had a midline crest on the top of
the skull. Their beaks often had toothless tips and they
may have been predators of molluscs and crustaceans.
Finally, the strangest pterodactyloids must be the
azhdarchoids. These include the long-necked azhdarchids and the crested tapejarids. Azhdarchids
may have exceeded wingspans of 11 m and were
widely distributed in the Late Cretaceous. They may
have been ecological generalists akin to storks, and
were probably not specialist carrion feeders or mud
probers as has been proposed. Determining the life
style of the tapejarids is more difficult. The vaguely
parrot-like skull of Tapejara (Figure 6) from Lower
Cretaceous Brazil led some workers to propose that
it was a fruit eater, but it might better be imagined
as an auk analogue. Recently, it has been suggested

that Thalassodromeus, also from Lower Cretaceous

Brazil, was a fish eater that trawled its blade-like
lower jaw through the water.
Pterosaur Palaeobiology
Because pterosaurs are unique and extinct, reconstructing their palaeobiology is difficult and nothing
is known about several aspects of their lives. Limited
evidence does allow us, however, to reconstruct their
sensory abilities, feeding behaviours, and styles of
locomotion.
The large orbits of pterosaurs show that they had
large eyes, and the abundance of visual display
phylum Mollusca, is one of the most familiar of all
invertebrate taxa. Modern representatives, such as
mussels, cockles, oysters, and scallops, are well
known from excursions to the coast, and in many
parts of the world they are important commercial
species. Their generally excellent fossil record has
allowed their evolutionary history to be traced back
to the Early Palaeozoic and, for much of this time,
they have been important components of many
faunas. From rather modest beginnings, they have
conquered a range of habitats from the deep sea to
freshwater, exploited a wide range of life habits
(from deep burrowing to swimming), and undergone
a near-exponential taxonomic proliferation, a spectacular example of an adaptive radiation.
Bivalves come in all manner of shapes and sizes,
from tiny, thin-shelled commensals that live in association with sea anemones, to giant clams and the
extinct rudists and inoceramids which reach(ed)
sizes well over 1 m. Shell morphology is extremely
plastic, but all are modifications of the same basic
theme. The intimacy of the shell morphology to life
habit has been a great benefit in reconstructing the life
habits of extinct bivalves, but has also frustrated
many attempts to establish the relationships between
different groups within the class. Bivalves have been
proven to be good palaeoenvironmental indicators,
but they have only limited use in biostratigraphy.
Freshwater mussels have been used to date fluvial
deposits in the Carboniferous Coal Measures of
Western Europe, and inoceramids have been used
for Late Cretaceous deep marine settings (e.g., in
New Zealand). In general, however, species of the370 FOSSIL INVERTEBRATES/Bivalv
es
class are too long lived and too facies specific to be of
any great value.
General Morphology
As the name implies, bivalves comprise two calcareous valves. These are arranged laterally (left and
right), are joined dorsally by a partially calcified elastic ligament, and enclose the soft tissue. Each valve
has clearly differentiated posterior and anterior features, i.e., inequilateral. The primitive arrangement,
retained by most bivalves, was to have a plane of
symmetry parallel to the commissure (the join between the two valves), resulting in valves which are
mirror images of one another (i.e., equivalve). Although this symmetry is found in virtually all bivalves

which live with the commissural valve perpendicular

to the substrate surface (orthothetic), it has been lost
in those which have adopted a pleurothetic habit
where they lie on one valve (e.g., oysters, scallops).
In these cases, there is a tendency for the two valves to
become dissimilar (i.e., inequivalve), typically with
the underlying valve becoming more bowl-like and
the upper one more reduced like a lid.
Shell Morphology
All bivalves possess a pair of shells which may be
shut to provide protection from both environmental
stresses (e.g., desiccation in the intertidal habitat) and
the threat of predation. Most shells are reasonably
robust, which has provided the class with a generally
excellent fossil record. Although shell morphology in
bivalves is very variable and intimately linked to their
life habits (see below), all shells are simple modifications of the basic shell secretion model used by all
shelled molluscs. The shell is secreted by the mantle
lobes and grows by marginal accretion, as evidenced
by the growth lines on the surface of the valve
(Figures 1A and 2A). These growth lines are particularly marked in bivalves from intertidal and shallow
temperate habitats, where the animals experience
pronounced seasonality and largely stop growing
during the winter months. Bivalves which experience
more equable conditions do not show such obvious or
regular patterns. Inspection of the growth lines in
sectioned valves shows that, although most shell material is added ventrally, the shell is also thickened
during growth (Figure 2B), demonstrating that the
entire mantle surface is responsible for adding material. The outermost part of the shell is an organic layer
called the periostracum secreted at the mantle edge
(Figure 2C). The thickness of the periostracum varies
between taxa, from less than 1 mm in oysters and
Figure 1 (A) Mercenaria mercenaria , a shallow burrowing bi
valve from the Pliocene of Florida. Note the prominent annual
growth bands. (B) Pecten maxima , a free living epifaunal scallop
from the Holocene Atlantic.
scallops to several hundred micrometres in some
mussels. In many cases, the periostracum is lost by
abrasion and decay during the life of the animal,
particularly on the older parts of the shell, and there
is no real prospect of it being preserved in any but the
most exceptional circumstances. The primary function of the periostracum is to act as the template on
which the calcareous part of the shell is deposited, but
it may also provide protection from both corrosive
waters and predators that dissolve the shell. It is
particularly noticeable that freshwater bivalves have
very thick periostraca.
The main part of the shell, however, is calcareous.
It is in effect a ceramic made up of calcium carbonate
crystals in an organic matrix (the latter accounting for
<5% of the dry weight of the shell). The proteinaceous matrix controls both the polymorph of calcium carbonate used and the arrangement of the
crystals. All bivalves contain aragonite in their shells
and the vast majority are wholly so. Some taxa, how-

ever, chiefly those exploiting epifaunal life habits, also

secrete calcite in their outer layers. The oysters have
taken this to its extreme and the bulk of the shell isFOSSIL INVERTEBRATES/Bival
ves 371
Figure 2 Details of shell formation in a generalized bivalve.
(A) Marked comarginal growth lines on the shell surface.
(B) Section through the shell along the line indicated in (A) show
ing the arrangement of growth lines within the shell. (C) The
relationship between the shell and the underlying mantle edge
(close up details of the circled area in (B)). i, inner mantle fold; m,
middle mantle fold; o, outer mantle fold.
calcitic, with aragonite being confined to the sites of
muscle attachment and the ligament.
Molluscan shell is immensely strong, in fact often
much stronger than vertebrate bone. There are a
number of different microstructures (Figure 3), each
with different mechanical properties, and most shells
are made up of two or three arranged in different
layers. Different taxa show different arrangements
and these are considered to be of phylogenetic significance. It is apparent that the earliest bivalves were
wholly aragonitic and chiefly composed of nacre
(Figure 3A), and that subsequent evolution has produced the wide array of microstructural arrangements
seen today. The effect of differing crystal sizes, amount
of organic material, and polymorph used has affected
the preservation potential of different taxa; many of
the Palaeozoic and Mesozoic taxa that were originally
aragonitic are either preserved as internal moulds or
are replaced by calcite.
Details of the internal features of the shell are
shown in Figure 4. The hinge plate is situated dorsally
and houses the ligament and teeth. The ligament is an
elastic, partially calcified layer that provides a very
energy-efficient opening mechanism. During valve
closure, energy is stored in the ligament as it is flexed
by the contracted adductor muscle(s) (Figure 4C).
When the muscle is relaxed, the ligament springs the
valves apart causing them to gape. This passive valve
opening mechanism is the reason why many fossil
bivalves are found in a disarticulated state. Although
the ligament itself is seldom preserved, its position
may be inferred from the presence of the ligament
pits in which it is anchored (Figures 4 and 5). Most
bivalves have teeth on the hinge plate which fit into
corresponding sockets on the opposite valve and function to keep the valves in perfect alignment. Both
ligamenture and dentition vary markedly amongst
higher taxa of bivalves, and both are often used as
informative characters in establishing phylogenies.
Some of the range of hinge plate architectures is
shown in Figure 5.
A number of attachment scars mark the locations
where muscles are anchored to the shell. The most
significant of these are the adductor scars (Figures 4A
and 4B). If the adductor scars are paired (i.e., dimyarian), they occur posteriorly and anteriorly. If an
animal is monomyarian, the single muscle (the posterior) occupies a more central position. In many
taxa, there is a thin pallial line running around the

shell a small distance from the ventral edges that

marks the attachment of the mantle to the shell. In
infaunal taxa, where the posterior mantle has been
fused and elongated to form siphons, the pallial line is
inflected forming the pallial sinus. The sinus represents the space into which the siphons are withdrawn
when the valves are shut. Other muscle attachment
scars may be more or less apparent, including the
insertions of the pedal musculature (particularly in
burrowers and byssate taxa).
Soft Part Anatomy
Bivalves are laterally compressed and, unlike most
molluscs, there is no head or radula. The internal
organs are enclosed by the two mantle lobes that are
joined dorsally (Figure 4C). The chief function of the
mantle is to secrete the shells, but the ventral edges of
each mantle lobe are differentiated into three folds
(Figure 2C), only the outermost of which is directly
concerned with shell manufacture. The innermost
fold controls water flow into and out of the mantle
cavity, whilst the middle fold has sensory capability.
In several bivalve groups (such as scallops), the
middle fold is well developed with tentacles and
eyes. In some taxa, the mantle is extended posteriorly
and fused to form a pair of siphons through which
water is directed into (inhalant) and out of (exhalant)
the mantle cavity.372 FOSSIL INVERTEBRATES/Bivalves
Figure 3 Scanning electron micrographs of four of the most common bivalve shell
microstructures. (A) Aragonitic nacre (from the
inner shell layer of Pinna nobilis ). (B) Aragonitic crossed lamellar structure
(from the outer layers of Corbula gibba ). (C) Foliated calcite
(from Ostrea edulis ). (D) Calcitic prisms (from the outer shell layer of Pinna
nobilis ). Scale bars for (A) (C) represent 10 mm and for
(D) represents 100 mm.
The mantle cavity is spanned by one or two adductor
muscles that attach to the shell and act antagonistically
with the ligament to close the valves on contraction
(Figure 4C). A significant part of the mantle cavity is
occupied by a pair of gills (the ctenidia) lying on either
side of the rest of the viscera. In most bivalves, the gills
are involved with both respiration and ciliary suspension feeding (filtering small particles out of the water
which are then transferred to the mouth by a pair of
labial palps). Recent bivalves show a number of different gill morphologies depending largely on the feeding process employed. Deposit feeders, e.g., Nucula,
have less well-developed (protobranch) gills, whilst
members of the Lucinidae augment their filter feeding
by energy gained from the activities of sulphide-oxidizing chemosymbiotic bacteria living within the modified
gills. The carnivorous septibranch bivalves (e.g.,
Cuspidaria, Poromya) suck in small prey (such as
amphipods) using their modified siphons. These extraordinary bivalves have lost their gills and respire
over the inner surface of the mantle. Other significant
organs within the mantle cavity include the gut, heart,
circulatory system, and the foot. The gut runs between
the anteriorly positioned mouth and the posterior anus,
and includes a complex stomach which, again, has a
number of configurations depending on the feeding

biology of the animal. Blood is circulated throughout

the animal by a three-chambered heart. A muscular
foot is present in all juvenile and most adult bivalves
and occupies the centre of the mantle cavity.
Naturally, the soft part anatomy of bivalves is very
seldom preserved, although preservation of gill and
muscle material has been reported in exceptional circumstances. Various details, however, can be inferred
from the study of the internal surface of the shells.
Apart from adductor muscle scars, the practised eye
may pick out the attachment points of more minor
muscles and impressions of radial muscles and blood
vessels within the mantle.
Ecology
Modern bivalves exploit a wide range of life habits.
Many burrow to varying depths within soft sediments,FOSSIL INVERTEBRATES/Bivalve
s 373
Figure 4 Various aspects of the internal morphology of bivalve
shells. (A) The left valve of a generalized burrowing dimyarian.
(B) The right valve of a generalized byssate scallop. (C) The rela
tionship between the two valves and their associated mantle
lobes, adductor musculature and ligament.
but others attach to or bore into hard substrates by a
variety of means; others have become free living, some
with the ability to swim. Most bivalves are marine,
exploiting niches from the intertidal zone down into
the abyssal depths, but successful groups (including
the oysters) have invaded more brackish conditions
and even freshwater, where modern unionid mussels
cause enormous damage as biofoulers. It is clear that
the most primitive bivalves were marine shallow
burrowers and that other life styles evolved later. It is
also apparent that many of the more specialized life
habits have evolved separately in a number of different lineages (i.e., polyphyletically). Seminal work by
S. M. Stanley firmly established how different aspects
of the morphology of living bivalves could be related
to their life habits, such that it is possible to use these
characteristics of extinct taxa to reconstruct the life
habits of fossil groups.
Burrowing
A large proportion of all bivalves (around 50% of all
modern families) burrow into soft sediments using the
foot. Most are equivalve and are isomyarian (i.e., the
posterior and anterior adductor muscles are of equal
size). The depth to which they burrow varies between
taxa, from those which lie just under the surface with
the edge of the shell virtually level with the sediment
water interface (e.g., Cerastoderma; Figure 6C), to
depths of several centimetres (e.g., Mya; Figure 6B),
with Panopea reaching spectacular depths of up to
1 m. The key to successful burrowing is maintaining
contact with the seawater in order to continue
both feeding and respiration. This is achieved by
the siphons, snorkel-like extensions of the posterior
mantle. The length of the siphons, and therefore the
depth of burrowing, can be inferred from the shells by
the size of the pallial sinus; deeper burrowers have
more indented pallial sinuses, whereas very shallow
burrowers have no sinus at all (see Figure 6). Very

deep burrowers, such as Mya, have siphons so long

that they are unable to withdraw them fully into the
shell when it shuts, and have a permanent posterior
siphonal gape through which they protrude. Shallow
burrowers generally have strong, robust shells, often
with a pronounced radial or concentric ornament that
may assist the burrowing process or help the animal
remain locked into the sediment. Deeper burrowers
tend to have thinner shells and are often smooth
shelled. Although the foot is never preserved, its presence may be inferred from the pedal musculature on
the inside of the valves and, in cases where the animal
is a rapid and deep burrower (such as the razor shell
Ensis), the foot may be so well developed as to require
an anterior pedal gape.
It is clear from studies of the siphons of living
bivalves that they are constructed in a number of different ways, suggesting that the deep burrowing habit
has evolved independently in several clades.
Attachment
Almost all larval bivalves attach to the substrate, if
only briefly, with tanned protein threads (the byssus)
secreted by a gland at the base of the foot. In a large
number of taxa, this habit has been neotenously
retained into adulthood, and again it is clear that
this has happened repeatedly in different groups.
Byssate bivalves fall into two categories: those like
Pinna (Figure 6A) and Modiolus that are orthothetic
and live attached to clasts within the sediment in
which they are partially buried (endobyssate), and
those that are attached to the surface of hard
substrates (epibyssate), either in an orthothetic (e.g.,
Mytilus; Figure 7D) or a pleurothetic (e.g., Isognomon ephippium; Figure 7C) orientation. Orthothetic
byssate bivalves tend to be equivalve and have much
reduced anteriors. This anterior reduction is reflected374 FOSSIL INVERTEBRATES/
Bivalves
Figure 5 Selected hinge plates showing some of the variety of ligament insertion
and arrangement of teeth. (A) Cerastoderma edule :
heterodont dentition with two centrally placed cardinal teeth and two lateral te
eth. (B) Venus casina : heterodont (similar to Cerastoderma
but with no lateral teeth). (C) Arca tetragona : taxodont dentition with numerou
s teeth arranged in a row; the ligament forms a chevron
pattern on the broad triangular area below the umbones. (D) Chlamys varia : two
simple teeth with the internal ligament occupying a
triangular pit below the umbones.
Figure 6 Morphology and mode of life of bivalves living in or partially within s
oft substrates. (A) Pinna nobilis . Not to scale. (B) Mya
truncata . (C) Cerastoderma edula .
in the adductor musculature, which (although still
dimyarian) is heteromyarian, with the anterior adductor much smaller than the posterior (Figure 7D).
Pleurothetic byssate bivalves are often markedly
inequivalve, with the lower valve (which in the majority of cases is the right) often larger than the other.
Although they are dimyarian early in ontogeny, the
adults are monomyarian, having lost the anterior
muscle during ontogeny; the remaining posterior
muscle is often large and centrally placed (Figure 7C).
The presence of a byssus may be inferred from either a

slight gape between the valves through which it passes

(the byssal gape), or more obviously the byssal notch in
scallops (Figure 4B).FOSSIL INVERTEBRATES/Bivalves 375
Figure 7 Morphology and mode of life of bivalves living on or boring into substr
ates. (A) Spengleria rostrata . (B) Spondylus americanus .
(C) Isognomon ephippium . (D) Mytilus edulis . (E) Placuna placenta . (F) Grypha
ea arcuata . Not to scale.
Whereas byssate attachment is flexible and also
renewable, some bivalves permanently attach to
hard substrata by a calcareous cement. The cemented
habit always succeeds a byssate phase and it is clear
that it has evolved independently in a number of different clades (e.g., oysters, rudists, and a number of
pectinoids including Spondylus). Cemented bivalves
are often easily recognizable from their irregular
morphology, developed because of the requirement
to conform to substratal irregularities, and are markedly inequivalve. As they tended to evolve from
pleurothetic byssate stock, most are also monomyarian (Figure 7B). Most cement mainly by the right
valve, but a major group, the oysters, do so by the
left valve. The size of the attachment scar varies and
the substrate may be instantly recognizable; for
example, oysters were often attached to ammonite
shells, even if the scar and substrate are no longer
attached. Most cementers have thick, robust shells
and may be extravagantly ornamented with spines or
flanges (e.g., Spondylus; Figure 7B).
Free Living
A number of different taxa have independently abandoned attachment to become free living on softer
sediments. Here, the challenge is not to sink into the
substrate, and this has been solved in two ways. The
first is by adopting a snow-shoe-type morphology,
i.e., resting on a large surface area, epitomized by the
wafer-thin window pane shell Placuna (Figure 7E).
Alternatively, they may be semi-submerged in the soft
sediment like an iceberg. This strategy is inferred for
the thick-shelled devils toenail Gryphaea, common in
Mesozoic clay facies (Figure 7F). These aberrant
oysters clearly had a cemented phase, marked by a
small attachment scar at the umbo. A few free-living
bivalves (notably several groups of scallops) also
have the ability to swim short distances if they are
threatened. These have smooth hydrodynamic shells
and a well-developed posterior adductor muscle
whose vigorous contraction provides the propulsion.
Boring
A number of groups, once again polyphyletically,
have evolved to excavate burrows in hard substrates
by boring. The most successful of these, the mytilid
lithophagids, do so principally by acidic secretions
(which presents its own challenge of not dissolving
its own shell), whilst others, e.g., Pholas, bore at least
partly by physically rasping the substrate with small
projections on the outside of the shell. Some of the
most bizarre borers are the teredinids which excavate
long cylindrical boreholes in wood and have the enzymatic capability of digesting the cellulose. These
shipworms are thought to have been part of the

undoing of the ships of the Spanish Armada.

Members of the boring group as a whole have a
very varied morphology, but may be easily recognized
because they are almost invariably fossilized within
Statesman of London, called it "difficult, though not absolutely so." The Times
of
London, did an informal poll, asking seven reporters and a math student to read
it and
report on its accessibility. The verdict was mixed. "It all made beautiful sense
as I read it,
though it tended to vanish like a dream when I put the book down," one wrote.
Albert Einstein once said that scientific theories should be able to be describe
d so simply
that a child could understand them. Complaints that modern physics fails this st
andard
abysmally are as old, well, as modern physics, and are not confined to the child
like
public.
The story goes that when the astronomer Arthur Eddington, whose observations of
light
bending during a solar eclipse in 1919 confirmed Einstein s general theory of re
lativity,
was congratulated by a colleague on being one of the three people in the world w
ho
understood the abstruse theory, Eddington fell uncharacteristically silent. Chid
ed for
exhibiting a false modesty, Eddington replied, on the contrary, that he had been
trying to
imagine who the third person could be.
This newspaper s early accounts of Einstein s and Eddington s 1919 breakthroughs
focused on the theory s incomprehensibility. "Efforts made to put in words intel
ligible to
the nonscientific public the Einstein theory of light proved by the eclipse expe
dition so
far have not been very successful," began a article on Nov. 10, 1919.
Niels Bohr, one of the founders of quantum mechanics, once said that anyone who
was
not outraged on hearing about the theory -- with its waves acting as particles,
particles
acting like waves, and the microscopic randomness and uncertainty it ascribed to
nature - had not really understood it.
Recent advances have made it even harder to explain the universe. The latest ver
sion of
the putative theory of everything posits a universe with 10 or 11 dimensions, in
stead of
35the 3 of space and 1 of time of everyday experience, inhabited by wriggling st
rings or
membranes. Nevertheless, scientists go on gamely trying to tell us what they are
up to, in
a book-writing tradition that includes Darwin s "Origin of Species," and Einstei
n s,
"Relativity: The Special and the General Theory," written in 1916 and never out
of print.
Part of the lure of these books is the chance to reclaim one s citizenship in a
troubled and
baffling cosmos by hearing the word from the horse s mouth, from someone who has
touched the cosmic mystery personally. But another part is surely being treated
like an
adult, of entering a rough-hewn colleagueship by being trusted to put work into

deciphering statements like the one at the beginning of this essay, or to deal w
ith straight
talk of the nature of science and the universe.
Here, for example, is Dr. Hawking about those troublesome extra dimensions requi
red by
string theory but apparently unavailable for parking cars. "I must say that pers
onally, I
have been reluctant to believe in extra dimensions," he writes on Page 54 of the
new
book. "But as I am a positivist, the question Do extra dimensions really exist?
has no
meaning. All one can ask is whether mathematical models with extra dimensions pr
ovide
a good description of the universe."
In other words, if the experiments come out right, it doesn t matter. This could
be
considered jarring if you cling to the notion that science is the search for a r
eality that is
deeper than the measurements on a laboratory table. But, quantum theory and rela
tivity
have taught us, science is about what can be observed and measured or it is abou
t nothing
at all. In science, as in democracy, there is no hidden secret knowledge, all th
at counts is
on the table, observable and falsifiable. All else is metaphysics.
When it comes to putting the goods on the table without condescending, Dr. Hawki
ng is a
genius. While many authors of science books plough through chapters full of
fundamentals before getting to the new stuff, Dr. Hawking, with perhaps a height
ened
appreciation of time, breezes speedily to the frontier without apologies.
For those who cannot keep up, Dr. Hawking has also provided a legacy. The succes
s of
his earlier book and that of Carl Sagan s "Cosmos" are widely credited with havi
ng given
a commercial lift to the science-book genre, helping pave the way for efforts li
ke "The
Elegant Universe," by Dr. Brian Greene, a Columbia University string theorist; "
The
Inflationary Universe," by Dr. Alan Guth, cosmologist at the Massachusetts Insti
tute of
Technology; and "The Quark and the Jaguar," by the Nobel laureate Murray Gell-Ma
nn.
To the extent that Dr. Hawking s earlier success has spawned imitators and widen
ed the
circle of readers and their sophistication, he has engineered a kind of positive
feedback,
and he has increased the odds that the readers will follow him and get to the en
d of the
book this time.
Copyright 2002 The New York Times Company
36Mysteries of the Universe
ENDLESS POSSIBILITIES
The End of Everything
By DENNIS OVERBYE
In the decades that astronomers have debated the fate of the expanding universe
-whether it will all end one day in a big crunch, or whether the galaxies will sa
il apart
forever -- aficionados of eternal expansion have always been braced by its seemi

ngly
endless possibilities for development and evolution. As the Yale cosmologist Dr.
Beatrice
Tinsley once wrote, "I think I am tied to the idea of expanding forever."
Life and intelligence could sustain themselves indefinitely in such a universe,
even as the
stars winked out and the galaxies were all swallowed by black holes, Dr. Freeman
Dyson,
a physicist at the Institute for Advanced Study, argued in a landmark paper in 1
979. "If
my view of the future is correct," he wrote, "it means that the world of physics
and
astronomy is also inexhaustible; no matter how far we go into the future, there
will
always be new things happening, new information coming in, new worlds to explore
, a
constantly expanding domain of life, consciousness, and memory."
Now, however, even Dr. Dyson admits that all bets are off. If recent astronomica
l
Transcription Control in
Eukaryotes
Transcription in eukaryotes differs from that in
prokaryotes in two main respects. In eukaryotes, one gene codes for a single polypeptide
(monocistronic transcription unit) and the initial transcript is processed into mature messenger mRNA. This involves intron splicing (see
p. 50) and substantial modification of the ends
of the primary transcript.
A. Prototype of a eukaryotic structural
gene
A structural gene is a gene that codes for a polypeptide gene product. It can be divided into sections involved in transcription (transcription
unit) and regulatory sequences. Regulatory
sequences are located both upstream (the 5!
direction) and downstream (the 3! direction) of
the gene. In addition, internal regulatory
sequences may occur in introns. Some regulatory sequences are located far from the gene.
Together with the promoter (see p. 206), they
are required to regulate transcription.
nosine is methylated in position 7, as are the
two initial ribose residues at the beginning of
the RNA chain. Except for the mRNAs transcribed by DNA viruses, eukaryotic mRNA usually contains a single protein-coding sequence
(monocistronic messenger).
D. Polyadenylation at the 3! end
Eukaryotic termination signals have been less
well recognized than the regulators of gene activity at the 5! end. Eukaryotic primary transcripts are split by a specific endonuclease
shortly after the sequence AAAUAA. Subsequently, about 100 250 adenine nucleotides
are attached to the 3! end of the transcript by
means of a poly(A)-polymerase (polyadenylation). The poly(A) end binds to a protein. All
mRNAs, except those that code for histone proteins, possess a poly(A) terminus.

(Figures after Singer and Berg, 1991).

References
Singer, M., Berg, P.: Genes & Genomes. Blackwell
Scientific, Oxford, 1991.
B. Prototype of mature eukaryotic
mRNA
Mature eukaryotic mRNA is produced from its
precursor RNA by the removal of introns, addition of a 5! cap at the 5! end, and addition of
numerous adenine nucleotides at the 3! end
(polyadenylation). A noncoding sequence (5!
leader) is located in front of the translation start
signal (AUG), and a trailer sequence, at the 3!
end in back of the translation stop signal (UAA).
Both addition of the 5! cap and polyadenylation
involve enzymatic reactions.
C. 7-Methyl-guanosine cap
The translation of eukaryotic mRNA is similar to
that of prokaryotic mRNA, with two distinct
differences: (1) transcription and translation
occur at different locations in the eukaryotic
cell: transcription occurs in the cell nucleus,
and translation in the cytoplasm; (2) the 5! and
3! ends of eukaryotic mRNA have special structures. The structure at the 5! end is called a cap.
Through the action of guanosine-7-methyltransferase, guanosine is bound by a triphosphate bridge to the first and second ribose
groups of the precursor mRNA chain. The guaPassarge, Color Atlas of Genetics 2001 Thieme
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ion Control in Eukaryotes
Passarge, Color Atlas of Genetics 2001 Thieme
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215216
Fundamentals
Regulation of Gene Expression
in Eukaryotes
Precisely regulated gene expression is a prerequisite for producing and maintaining the
many different types of cells and tissues of a
multicellular organism. Cells differentiate into
their particular cell types by means of combinations of expressed and repressed genes. During
differentiation the tightly regulated genes function in the order, usually sequential, required
for a particular cell fate (developmental pathways). Many regulator genes and their proteins
have been identified (cf. part III, Genetics and
Medicine). The following outlines some important principles of the specific control of gene expression in eukaryotic cells.
A. Levels of control of eukaryotic gene
expression
In principle, expression can be regulated at four
distinct levels. The first and by far the most important is primary control of transcription.
Processing to mature RNA can be regulated at
the level of the primary RNA transcript. A
frequently observed process is alternative splic-

ing (see D). Translation can be varied by RNA

editing (see B for an example). At the protein
level, posttranslational modifications can determine the activity of a protein. The cleavage of
preproinsulin to form mature insulin, glycosylation or hydroxylation, and protein folding are
some examples (see p. 32, 362).
B. RNA editing
RNA editing modifies genetic information at the
RNA level. An important example is the apolipoprotein-B gene involved in lipid metabolism. It
encodes a protein of 4538 amino acids, apolipoprotein B. This is synthesized in the liver and
secreted into the blood, where it transports
lipids. A related shorter form of the protein with
2153 amino acids, Apo B-48 (250 kDa, instead of
512 kDa for Apo B-100), is synthesized in the intestine. An intestinal deaminase converts a cytosine in codon 2158, CAA (glutamine), to uracil
(UAA). This change results in a stop codon (UAA)
and thereby terminates translation at this site.
C. Long-range gene activation by an
enhancer
Enhancers control gene activity at a distance.
An enhancer is a distant site involved in initiation of transcription (see p. 206). It may be located either upstream or downstream of the
same DNA strand (cis-acting) or on a different
DNA strand (trans-acting). Enhancer elements
provide tissue-specific or time-dependent regulation. It is unclear how enhancers can exert
their effect from a considerable distance. One
model suggests that DNA forms a loop between
enhancer and promoter. Activator proteins
bound to the enhancer, e.g., a steroid hormone,
could then come into contact with the general
transcription factor complex at the promoter.
Others might function as repressors (cf. transcription control in prokaryotes, p. 210).
D. Alternative RNA Splicing
A DNA segment can code for different forms of
mRNA when different introns are removed from
the primary transcript (alternative splicing). By
means of alternative gene splicing, a gene can
code for different, albeit similar gene products.
This allows a high degree of functional flexibility. Numerous examples of differential RNA
splicing are known for mammalian genes. For
example, the primary transcript for the calcitonin gene contains six exons. They are spliced
into two different types of mature mRNA. One,
consisting of exons 1 4 (but not exons 5 and 6),
is produced in the thyroid and codes for calcitonin. The other consists of exons 1, 2, 3, 5, and
6, but not exon 4. It codes for a calcitonin-like
protein in the hypothalamus (calcitonin generelated product, CGRP).
References
Alberts, B., et al: Essential Cell Biology. An Introduction to the Molecular Biology of the Cell.

Garland Publishing Co., New York, 1998.

Lewin, B.: Genes VII, Oxford Univ. Press, Oxford,
2000.
Blackwood E.M., Kadonga J.F.: Going the distance: A current view of enhancer action.
Science 281 :60 63, 1998.
Stryer, L.: Biochemistry. 4 th ed. W.H. Freeman &
Co, New York, 1995.
Watson J.D., et al.: Molecular Biology of the
Gene. 4 th ed. Benjamin/Cummings Publishing Co., Menlo Park, California, 1987.
Passarge, Color Atlas of Genetics 2001 Thieme
All rights reserved. Usage subject to terms and conditions of license.217
Regulation of Gene Expression in Eukaryotes
1
Apo B-100
Glu
CAA
Cytosol
5
mRNA unedited
Nucleus
DNA
UAA
5
3
Cytosine deamination
by intestinal deaminase
3
2158
Apo B-84
B. RNA editing
Activator protein
Control of
processing
(alternative
splicing)
Transcription
start site
5
3
Enhancer
mRNA
Promoter
Binding of an
activator protein
to the transcription
complex
Control of
translation
(mRNA editing)
Enhancer
Protein
Activator
protein
Transcription factors
Control of
protein activity
active
Translation

stop
1
Control of
transcription
Primary transcript
4538
inactive
A. Levels of control of eukaryotic
gene expression
Transcription
Promoter with
transcription
factors and
RNA polymerase II
C. Long-range gene activation by an enhancer
Calcitonin gene
5
Exon 1
Exon 2
Exon 3
Primary RNA transcript
5
1
mRNA
5
2
3
C cells in thyroid
1
2
Exon 4 Exon 5
4 5
Exon 6 3
6 3
Transcription
RNA processing
3
4
3
5
Translation
Calcitonin
Hypothalamus
1
2
3
5
6
Translation
Different
gene products
D. Alternative RNA splicing
Passarge, Color Atlas of Genetics 2001 Thieme
All rights reserved. Usage subject to terms and conditions of license.
CGRP
(Calcitonin
gene-related
peptide)
3 218
Fundamentals

DNA-Binding Proteins
Regulatory DNA sequences interact with proteins to exert proper functional control. Regulatory proteins can recognize specific DNA
sequences because the surface of the proteins
fits precisely onto the DNA surface. Three basic
groups of regulatory DNA sequences can be distinguished: (1) sequences that establish the
exact beginning of translation; (2) DNA segments that regulate the end, or termination;
and (3) DNA sequences near the promoter that
have specific effects on gene activity (repressors, activators, enhancers, and others).
A. Binding of a regulatory protein to
DNA
Gene regulatory proteins can recognize DNA
sequence information without having to open
the hydrogen bonds within the helix. Each base
pair represents a distinctive pattern of hydrogen bond donors (example shown in red) and
hydrogen acceptors (example shown in green).
These proteins recognize the major groove of
DNA, where binding takes place. Here a single
contact of an asparagine (Asn) of a gene-regulatory protein with a DNA base adenine (A) is
shown. A typical area of surface-to-surface contact involves 10 20 such interactions. (Figure
redrawn from Alberts et al., 1998, p. 276).
B. An helix inserts into
m
jor
groove of oper
tor DNA
One p
rt of the protein,
n helix (the
sequence-re
ding or recognition helix) is inserted into the m
jor groove of DNA. Here the
sequence Q-Q-Q-S-T (glut
mine Q, serine S,
threonine T) in the recognition sequence of the
b
cterioph
ge 434 repressor bonds with
specific b
ses in
m
jor groove of oper
tor
DNA. (Figure redr
wn from Lodish et
l., 2000,
p. 351).
C. Zinc finger motif
Another group of proteins
re c
lled zinc fingers
bec
use they resemble fingers (see D). They
re
involved in import
nt functions during embryonic development
nd differenti
tion. The b
sic
zinc finger motif consists of
zinc
tom connected to four
mino
cids of
polypeptide
ch
in. Here, two histidine (H)
nd two cysteine
(C) residues
re shown in the schem
on the
left. The three-dimension
l structure on the
right consists of
n
ntip
r
llel sheet (amino
acids 1 10), an helix (
mino
cids 12 24),

nd the zinc connection. Four
mino
cids, cysteines 3
nd 6
nd histidines 19
nd 23,
re
bonded to the zinc
tom
nd hold the c
rboxy
(COOH) end of the helix to one end of the
sheet. (Figure redrawn from Alerts et al., 1994,
p. 411).
D. Zinc finger proteins ind to DNA
The interaction with DNA is strong and specific.
Each protein recognizes a specific DNA

sequence. As the numer of zinc fingers can e

varied, this type of DNA-inding has great evolutionary flexiility. (Figure redrawn from Alerts et al. 1994).
E. Binding to a response element
Many hormones and growth factors activate
cell-surface receptors. In contrast, steroid hormones enter the target cells and interact there
with a specific receptor protein in the cytosol.
The hormonereceptor complex then migrates
to specific sites of DNA. The hormone-inding
domain will prevent inding to DNA unless the
hormone is present. Activated receptors ind to
specific DNA sequences called hormone response elements (HREs). Each polypeptide
chain of the receptor contains a zinc atom
ound to four cysteines (1). The skeletal model
shows the two DNA-inding domains inding
to the HRE in two adjacent major grooves of the
target DNA (2). The space-filling model shows
how tightly the recognition helix of each dimer
of this protein fits into the major groove of DNA
shown in red and green (3). (Figure redrawn
from Stryer, 1995, p. 1002).
References
Alerts, B., et al.: Molecular Biology of the Cell.
3 rd ed. Garland Pulishing Co., New York,
1994.
Alerts, B., et al.: Essential Cell Biology. Garland
Pulishing Co., New York, 1998.
Lodish, H., et al.: Molecular Cell Biology. 4 th ed.
Scientific American Books, F.H. Freeman &
Co., New York, 2000.
Stryer, L.: Biochemistry. 4 th ed. W.H. Freeman &
Co., New York, 1995.
Tjian, R.: Molecular machines that control
genes. Sci. Am. 272 :38 45, 1995.
Passarge, Color Atlas of Genetics 2001 Thieme
All rights reserved. Usage suject to terms and conditions of license.DNA-Bindin
g Proteins
219
DNA-inding protein
Asn
Major groove
Donor
CH 3
T
H
CH 2
C
H
O N
N H
N
N
N
A
To sugar
25
N

R
K
V
H
Q
N
S
T
To sugar
H
Minor groove
A. Binding of a regulatory protein to DNA
HOOC
Q
H
N
O
Q
H Acceptor
H
HN
O
23
3
C
Zn
Q
K
1
Y
B. An ! helix inserts into a major
groove of operator DNA
25
NH 2
HOOC
His
23
C
L
6 C
H 19
R
E
S
L R
A S
His 19
F 10
S
K
V
Zn
E
12
12
D. A zinc finger protein inds to DNA
Cys
443
Cys
440

Zn
1.
Cys
457
2.
Cys
3
1
H 2 N
Zn
C. Zinc finger motif
Cys
460
Cys
6
Zn
3.
E. Binding to a response element
Passarge, Color Atlas of Genetics 2001 Thieme
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10220
Fundamentals
Other Transcription Activators
Transcription activators are dimeric proteins
with distinct functional domains: a DNA-inding domain and an activation domain. The DNAinding domain interacts with specific regulatory DNA sequences. The activation domain interacts with other proteins that stimulate transcription. Transcription activators participate in
the assemly of the initiation complex, for example, y stimulating the inding of transcription factor IID (TFIID, see p. 212) to the promoter. Other activators may interact with
general transcription factors. They provide a
second level of transcriptional control.
A. Leucine zipper dimer
Most DNA-inding regulatory proteins recognize specific sites as dimers. One part of the
molecule serves as the recognition molecule,
the other stailizes the structure. A particularly
striking example is given y proteins with a
leucine zipper motif. The name is derived from
the asic structure. Two helices
re joined like

zipper by periodic
lly repe
ted leucine residues loc
ted
t the interf
ce of the two helices.
The two helices sep
r
te, form
Y-sh
ped
structure,
nd extend into the m
jor groove of
the DNA (1). Leucine zipper proteins m
y be homodimers with identic
l subunits (2, 3) or heterodimers with different
lbeit simil
r subunits
(4). The
bility to form unlike dimers (heterodimeriz
tion) gre
tly exp
nds the spectrum of
specificites. The use of combin
tions of different proteins to control cellul
r functions is
c
lled combin
tori
l control. (Figure redr
wn
from Alberts et
l., 1994).
A DNA-binding motif rel
ted to the leucine zipper is the helixloophelix (HLH) motif (not
shown). The HLH motif consists of one short

helix
nd one longer helix. The two helices

re connected by
flexible loop of protein.
ment
re regul
ted by steroids (steroid-responsive tr
nscription). The l
tter include glucocorticoids
nd miner
locorticoids, the steroids of
glycogen
nd miner
l met
bolism; sex hormones, which function in embryonic sex differenti
tion
nd control of reproduction;
nd
others. Norm
l bone development
nd function

re under the control of steroidlike vit
min D.
Another steroidlike hormone is retinoic
cid,
n
import
nt regul
tor of differenti
tion during
embryogenesis (morphogen). These hormones
initi
te their physiologic
l effects by
ssoci
tion with corresponding steroid-specific tr
nscellul
r receptors (hormonereceptor complex).
C. Evidence of
protein-binding
region in DNA
Protein-binding regions in DNA represent regul
tory
re
s; thus, their
n
lysis c
n yield some
insights into gene regul
tion. Protein-binding
DNA regions c
n be demonstr
ted in sever
l
w
ys. With b
nd-shift
n
lysis (1), proteinbound
nd non-protein-bound DNA fr
gments

re differenti
ted using gel electrophoresis in
direction tow
rds the sm
ll fr
gments,
DNA
fr
gment th
t is p
rt of
DNA-protein complex
migr
tes more slowly th
n
free DNA fr
gment
of the s
me size. The DNA-protein complex is
found
t
different position (b
nd shift). DNA
footprinting (2) is
nother procedure for identifying protein-binding sites on DNA. The principle of DNA footprinting is th
t
proteinbound DNA region, e.g., the polymer
se-promoter complex, is protected from the effects of

DNA-cle
ving enzyme (DNA
se I). Previously
isol
ted DNA is cut into different fr
gments by
DNA
se,
nd the fr
gments
re sorted
ccording to size by gel electrophoresis. Since the DNA
protein-binding region is protected from cle
v
ge by DNA
se I (DNA
se I protection experiment), DNA b
nds from the binding region
re
missing (footprint).
B. Activ
tion by steroid hormone
binding References
Tr
nscription
l enh
ncers
re regul
tory regions of DNA th
t incre
se the r
te of tr
nscription. Their sp
cing
nd orient
tion v
ry rel
tive
to the st
rting point of tr
nscription. An enh
ncer is
ctiv
ted by binding to
hormonereceptor complex. This
ctiv
tes the promoter,

nd tr
nscription begins (
ctive gene). Numerous import
nt genes in m
mm
li
n develop- Alberts, B., et
l.: Molecul
r Biology
of the Cell.
3 rd ed. G
rl
nd Publishing Co., New York,
1994.
Alberts, B., et
l.: Essenti
l Cell Biology. G
rl
nd
Publishing, Co., New York, 1998.
Lodish, H., et
l.: Molecul
r Cell Biology. 4 th ed.

Scientific Americ
n Books, F.H. Freem
n &

Co., New York, 1999.
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.Other Tr
n
scription Activ
tors
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ss
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ge subject to terms
nd conditions of license.
221222
Fund
ment
ls
Inhibitors of Tr
nscription
nd
Tr
nsl
tion
A number of n
tur
l
nd
rtifici
l subst
nces

re
ble to inhibit tr
nscription or tr
nsl
tion.
They c
n be used to tre
t c
ncer or
s
ntibiotics
to tre
t infections. Although most subst
nces

re unspecific
nd not suit
ble for ther
peutic
purposes, some
re very specific
nd therefore
import
nt for the underst
nding of tr
nscription
nd tr
nsl
tion or for ther
py. B
sic
lly,
one c
n distinguish whether
n
gent interferes
with tr
nscription or with tr
nsl
tion.
A. Insertion of
ctinomycin D
between
GC b
se p
ir
Actinomycin D is
complex polypeptide produced by
species of streptomyces b
cteri
. It
consists of
phenox
zone ring with two symmetric
l side ch
ins (1). It
cts by becoming interc
l
ted between two neighboring GC b
se
p
irs in double-str
nded DNA. Viewed from the
side (2), the inserted
ctinomycin D molecule is
seen very distinctly within the DNA double
helix. In the view from
bove (3) the
ctinomycin D molecule forms
n
rrow l
yer within the
DNA double helix, bound by the two neighboring GC b
se p
irs. The degree of inhibition by

ctinomycin D v
ries gre
tly. High concentr
tions of
ctinomycin D block replic
tion,
where
s low concentr
tions suffice to inhibit
tr
nscription.
bosome,
nd the protein synthesis ends prem
turely.
(Figures A
nd B from Singer
nd Berg, 1991).
C. Inhibitors of protein synthesis
Numerous n
tur
lly occurring
nd
rtifici
lly
produced subst
nces inhibit protein synthesis
by inhibiting tr
nscription or cert
in ph
ses of
tr
nsl
tion. Some h
ve clinic
l signific
nce
s

ntibiotics; others
re toxicologic
lly signific
nt. An ex
mple for the specificity of some inhibitors is -
m
nitin,
dicyclic oct
peptide of
the fungus Am
nit
ph
lloides. In very low concentr
tions, it binds to RNA polymer
se II
nd
thereby blocks the form
tion of precursor
mRNA in euk
ryotes. In contr
st, RNA polymer
se I is insensitive to this toxin,
nd polymer
se III binds to it only in higher concentr
tions. (D
t

fter Singer
nd Berg, 1991).
References
Singer, M., Berg, P.: Genes & Genomes. Bl
ckwell
Scientific, Oxford, 1991.

B. Puromycin imit
tes
n
mino
cyl

tRNA
Puromycin,
polypeptide from Streptomyces
lboniger, blocks polypeptide synthesis in the ribosomes of prok
ryotes
nd euk
ryotes. Its
ction is b
sed on the structur
l simil
rity with
n

mino
cyl tRNA. An
mino
cyl tRNA is
tRNA
molecule with
n
mino
cid
tt
ched to its 3!
end. Norm
lly
peptide bond is formed by peptidyltr
nsfer
se between the
mino group of
the incoming
mino
cyl tRNA
t the A (
mino
cyl) position
nd the c
rboxyl group of the
peptidyl tRNA
t the P (peptidyl) position. The
structure of puromycin resembles th
t of

mino
cyl tRNA, but l
cking
n inter
ction with
the codon it c
nnot be
tt
ched to the A position in the ribosome. The resulting polypeptidylpuromycin
dduct is removed from the riP
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.Inhibitors
of Tr
nscription
nd Tr
nsl
tion
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.
223224
Fund
ment
ls
DNA Methyl
tion
Methyl
tion of cytosine residues in DNA pl
ys

n import
nt role in gene regul
tion. DNA
methyl
tion is required for norm
l embryonic
development. Genomic imprinting, X chromosome in
ctiv
tion, chrom
tin modific
tion,
nd
silencing of endogenous retroviruses
ll depend
on est
blishing
nd m
int
ining proper methyl
tion p
tterns. DNA methyl
tion is genespecific
nd occurs genome-wide. Two types of
methyltr
nsfer
se c
n be distinguished by their
b
sic functions: m
inten
nce methyl
tion
nd
de novo methyl
tion.
A. M
inten
nce methyl
tion
This type of methyl
tion is responsible for

dding methyl groups to the newly synthesized
DNA str
nd
fter replic
tion
nd cell division.
The methyl
ted sites in the p
rent
l DNA (1)

fter replic
tion (2) serve
s templ
te for correct methyl
tion of the two new str
nds. This
ensures th
t the previous methyl
tion p
ttern
is correctly m
int
ined (3). It results in both
d
ughter str
nds being methyl
ted
t the s
me
sites
s the p
rent
l DNA. The enzyme responsible for this is Dnmt1 (DNA methyl
se 1).
Its function is essenti
l. Mice deficient in this
enzyme die
s
result of genome-wide demethyl
tion.
B. Recognition of
methyl
ted DNA
segment
Cert
in restriction enzymes do not cle
ve DNA
when their recognition sequence is methyl
ted
(1). The enzyme Hp
II cle
ves DNA only when
its recognition sequence 5!-CCGG-3! is not
methyl
ted (2). MspI recognizes the s
me 5!-

CCGG-3! sequence irrespective of methyl
tion

nd cle
ves DNA
t this site every time. This
difference in cle
v
ge p
ttern results in DNA
fr
gments of different sizes serves to distinguish the methyl
tion p
ttern of the DNA.
C. DNA methyl
tion de novo
This is the second type of DNA methyl
tion.
Here methyl groups
re
dded
t new positions
of both str
nds of DNA, not just in the hemimethyl
ted str
nd
s in m
inten
nce methyl
tion shown in A. Two genes for different
methyltr
nsfer
ses with overl
pping functions
in glob
l remethyl
tion h
ve been identified recently: Dnmt3

nd Dnmt3b. Unmethyl
ted
DNA (1) is methyl
ted by their enzymes (2) in

site-specific
nd tissue-specific m
nner (3).
T
rgeted homozygous disruption of the mouse
Dnmt3

nd Dnmt3b genes results in severe
development
l defects. Double homozygous
mut
nts die before d
y 11.5 of the 21-d
y
embryonic development.
D. Hum
n DNMT3B gene
Mut
tions in the hum
n gene DNMT3B encoding type 3B de novo methyltr
nsfer
se c
use

distinctive dise
se c
lled ICF syndrome (immunodeficiency, centromeric chromosom
l inst
bility,
nd f
ci
l
nom
lies, McKusick c
t
log
no. 242860). The centromeres of chromosomes
1, 9
nd 16, where s
tellite DNA types 2
nd 3

re loc
ted,
re unst
ble. Cl
ssic
l s
tellite DNA
is grossly undermethyl
ted in
ll tissues. The
hum
n gene (1) consists of 23 exons sp
nning
47 kb of genomic DNA. Six exons
re subject to

ltern
tive splicing. The protein (2) consists of
845
mino
cids with five DNA methyltr
nsfer
se motifs (I, IV, V, IX, X) in the C-termin
l region. The
rrows point to six different mut
tions. The mut
tion
t position 809 (3),

ch
nge of A to G in codon 809, i.e., GAC (Asp) to
GGC (Gly), le
ds to the repl
cement of
sp
r
gine (Asn) by glycine (Gly). Both p
rents

re heterozygous for this mut
tion.
(Figure
d
pted form Xu et
l., 1999).
References
Bird, A: DNA methyl
tion de novo. Science
286 :2287 2288, 1999.
H
nsen, R. S. , et
l.: DNMT3B DNA methyltr
nsfer
se gene is mut
ted in the ICF immunodeficiency syndrome. Proc. N
t. Ac
d.
Sci. 96 :14 412 14 417, 1999.
Ok
no, M., et
l.: DNA Methyltr
nsfer
ses
Dnmt3

nd Dnmt3b
re essenti
l for de
novo methyl
tion
nd m
mm
li
n development. Cell 99 :247 257, 1999.
Reik, W., Kelsey, G., W
lter, J.: Dissecting de novo
methyl
tion. N
ture Genet. 23 : 380 382,
1999.
Robertson, K.D., Wolffe, A.P.: DNA methyl
tion
in he
lth
nd dise
se. N
ture Reviews 1 :11
19, 2000.

Xu, G., Bestor, T., et
l.: Chromosome inst
bility

nd immunodeficiency syndrome c
used
by mut
tions in
DNA methyltr
nsfer
se
gene. N
ture 402 :187 191, 1999.
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.DNA Methyl

tion
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.
225226
Fund
ment
ls
Genomic Imprinting
Genetic contributions from both the m
tern
l

nd the p
tern
l sets of chromosomes
re necess
ry for
m
mm
li
n zygote to develop norm
lly. The re
son lies in
p
rent-of-originspecific expression of cert
in genes. The
genome cont
ins defined regions where only
the m
tern
l gene copy is expressed,
nd not
the p
tern
l copy,
nd vice vers
. This
llelespecific gene expression, depending on the
p
rent
l origin, results from so-c
lled genomic
imprinting. Genomic imprinting h
s import
nt
implic
tions for hum
n genetic dise
se (see
p. 398).
A. The import
nce of two different
p
rent
l genomes
Different development
l results
re observed
when the fem
le pronucleus is removed from

mouse zygote (1) before the pronuclei h
ve
fused
nd is repl
ced either by
nother m
le
pronucleus (2) or by
control (i.e., the s
me
s
removed) (3), or when the m
le pronucleus is
removed
nd repl
ced by
fem
le pronucleus
(4) or
control. If the fem
le pronucleus is repl
ced by
m
le pronucleus (2), the zygote first

ppe
rs norm
l. However,
fter impl
nt
tion
should ensue, ne
rly
ll
ndrogenotes f
il to
complete preimpl
nt
tion (2). Those few th
t,
r
rely,
re
ch
postimpl
nt
tion
develop
completely
bnorm
lly. Such embryos do not
develop beyond the 12-somite st
ge.
When
m
le pronucleus is repl
ced by
fem
le
pronucleus (4),
gynogenetic zygote, which
differs m
rkedly from the
ndrogenote. Here,

bout 85% of gynogenotes re
ch norm
l preimpl
nt
tion development. But
lthough the embryo
t first develops f
irly well, the extr
embryonic membr
nes
re
bsent or underdeveloped,
nd the gynogenote dies before
re
ching the 40-somite st
ge. (Figure redr
wn
from S
pienz

nd H
ll, 1995).
B. Hum
n embryonic development
depends on the presence of

m
tern
l
nd
p
tern
l genome
A n
tur
lly occurring hum
n
ndrogenetic zygote is
hyd
tidiform mole (1). This is
n
bnor-

m
l pl
cent
l form
tion cont
ining two sets of

p
tern
l chromosomes
nd none from the
mother. No embryo develops
lthough impl
nt
tion t
kes pl
ce. The pl
cent
l tissues develop
m
ny cysts (2). When only m
tern
l chromosomes
re present,
n ov
ri
n ter
tom
with
m
ny different types of fet
l tissues develops
(3). No pl
cent
l tissue is present in this n
tur
lly occurring gynogenetic zygote. In triploidy,
rel
tively frequent glob
l chromosom
l leth
l hum
n disorder (see p. 402), extreme hypopl
si
of the pl
cent

nd fetus is
observed when the
ddition
l chromosom
l set
is of m
tern
l origin (4). (Photogr
phs kindly
provided by Professor Helg
Rehder, M
rburg.)
C. Genomic imprinting is est
blished in
e
rly embryonic development
The imprint p
ttern present in som
tic cells (1),
with one
llele
ctive only either, the m
tern
l
or the p
tern
l, prop
g
ted through
ll mitotic
divisions. However, in primordi
l germ cells the
imprint is er
sed (2). During g
metogenesis the
imprint is reset (3). In the m
le germline
ll
g
metes receive the p
tern
l imprint
nd in the
fem
le germline
ll g
metes receive the m
tern
l imprint. After fertiliz
tion, the correct imprint p
ttern is present in the zygote (4). It is
m
int
ined through
ll subsequent cell divisions under the control of
region
l imprinting
center (see p. 398).
References
B
rlow, D.P.: G
metic imprinting in m
mm
ls,
Science 270 :1610 1613, 1995.
Horsthemke, B.: Genomisches Imprinting beim
Menschen: Grundl
gen und klinische Relev
nz. Biospektrum 4 :23 26, 1998.
Morrison, I.M., Reeve, A.E.: C
t
logue of imprinted genes
nd p
rent-of-origin effects
in hum
ns
nd
nim
ls. Hum. Mol. Genet.
7 :1599 1609, 1998.
Reik, W., Sur
ni, A., eds.: Genomic Imprinting.
IRL Press
t Oxford University Press, Oxford,
1997.
S
pienz
, C., H
ll, J.G.: Genetic imprinting in
hum
n dise
se. pp. 437 458. In: The Met
bolic
nd Molecul
r B
ses of Inherited Dise
se. 7 th ed. C.R. Scriver, et
l., eds. McGr
wHill, New York, 1995.
Sur
ni, A.: Imprinting
nd the initi
tion of gene
silencing. Cell 93 :309 312, 1998.
Tilghm
n, S. M.: The sins of the f
thers
nd
mothers: Genomic imprinting in m
mm
li
n development. Cell 96 :185 193,
1999.
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
munic
tion between cells to
ssure growth,
differenti
tion, specific functions in different
types of cells,
nd proper response to extern
l
stimuli. Specific cellcell inter
ctions between
different types of cells h
ve evolved. A common

leitmotif is the specific binding of
n extr
cellul
r sign
ling molecule (lig
nd) to
specific receptor of the t
rget cell to trigger
specific
function
l response. The v
st v
riety of
molecules involved in the m
ny different types
of cells c
n be cl
ssified into f
milies of rel
ted
structure
nd function (see Lodish et
l., 2000;
Alberts et
l., 1994). Two
re
s
re selected
here: the m
in intr
cellul
r functions controlling growth
nd the receptor tyrosine kin
ses.
A. M
in intr
cellul
r functions
controlling growth
Growth f
ctors
re
l
rge group of different extr
cellul
r molecules th
t bind with high specificity to cell surf
ce receptors (1). Their binding
to the receptor (2)
ctiv
tes intr
cellul
r sign
l
tr
nsduction proteins (3). This initi
tes
c
sc
de of events resulting in
ctiv
tion of other
proteins (often by phosphoryl
tion) th
t
ct
s
second messengers (4). Hormones of different
types
re
heterogeneous cl
ss of sign
ling
molecules (5). They enter the cell either by diffusion through the pl
sm
membr
ne or by
binding to
cell surf
ce receptor (6). Some hormones require
n intr
nucle
r receptor (7).
Eventu
lly the sign
l c
sc
de results in
ctiv
tion or in
ctiv
tion of tr
nscription f
ctors (8).
Before tr
nscription
nd tr
nsl
tion ensue,
n
el
bor
te system of DNA d
m
ge recognition

nd rep
ir systems (9) m
ke sure th
t cell prolifer
tion is s
fe (cell cycle control, 10). In the
event th
t f
ults in DNA structure h
ve not been
rep
ired prior to replic
tion,
n import
nt
p
thw
y s
crifices the cell by
poptosis (cell
de
th, 11). (Figure
d
pted from Lodish et
l.,
2000.)
B. Receptor tyrosine kin
se f
mily
Like the G protein-coupled receptors (GPCRs,
see p. 268)
nd their effectors, the receptor tyrosine kin
ses (RTKs)
re
m
jor cl
ss of cell
surf
ce receptors. Their lig
nds
re soluble or
membr
ne-bound growth f
ctor proteins. RTK
sign
ling p
thw
ys involve
wide v
riety of
other functions. Mut
tions in RTKs m
y send

prolifer
tive sign
l even in the
bsence of

growth f
ctor, resulting in errors in embryonic
development
nd different
tion (congenit
l
m
lform
tion) or c
ncer. Of the more th
n
twenty different RTK f
milies, five ex
mples
re
selected here: the epiderm
l growth f
ctor receptor (EGFR); insulin receptor (IR); fibrobl
st
growth f
ctor receptor (FGFR) types 1, 2,
nd 3;
pl
telet-derived growth f
ctor (PDGFR);
nd
RET (re
rr
nged during tr
nsform
tion).
These receptors sh
re structur
l fe
tures,
lthough they differ in function. All h
ve
single
tr
nsmembr
ne dom
in
nd
n intr
cellul
r tyrosine kin
se dom
in of slightly v
ried size. The
extr
cellul
r dom
ins consist of evolution
rily
conserved motifs: cystein-rich regions, im-

munoglobulin (Ig)-like dom
ins, fibronectin repe
ts in the tyrosine kin
se with Ig
nd the EGF.
RTK mut
tions c
use
group of import
nt
hum
n dise
ses
nd m
lform
tion syndromes.
The phenotypes of the mut
tions differ
ccording to the p
rticul
r type of RTK involved
nd
the type of mut
tion.
References
Alberts, B., et
l.: Molecul
r Biology of the Cell.
3 rd ed. G
rl
nd Publishing Co., New York,
1994.
Cohen, M.M.: Fibrobl
st growth f
ctor receptor
mut
tions, pp. 77 94, In: M.M. Cohen Jr.,
R.E. M
cLe
n, eds., Cr
niosynostosis, Di
gnosis, Ev
lu
tion,
nd M
n
gement. 2 nd ed.
Oxford University Press, Oxford, 2000.
Lodish, H., et
l.: Molecul
r Cell Biology (with
n

nim
ted CD-ROM). 4 th ed. W.H. Freem
n &
Co., New York, 2000.
Muenke, M., et
l.: Fibrobl
st growth f
ctor receptorrel
ted skelet
l disorders: cr
niosynostosis
nd dw
rfism syndromes, pp.
1029 1038, In: J.L. J
meson, ed., Principles
of Molecul
r Medicine. Hum
n
Press, Totow
, New Jersey, 1998.
Mnke, M., Schell, U.: Fibrobl
st-growth-f
ctor
receptor mut
tions in hum
n skelet
l disorders. Trends Genet. 11 : 308 313, 1995.
Roberton, S. C., Tyn
n, J.A., Donoghue, D.J.: RTK
mut
tions
nd hum
n syndromes: when
good receptors turn b
d. Trends Genet.
16 : 265 271, 2000.
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.Intr
cellu
l
r Sign
l Tr
nsduction Systems
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.
265266
Genetics
nd Medicine
Types of Cell Surf
ce Receptors
Specific receptors on cell surf
ces (
nd in the
nucleus or cytosol) convey cell-to-cell sign
ls
into the cells
nd the function
l
nswers. The
b
sic structures of their genes
re simil
r bec
use they h
ve been derived from
rel
tively
sm
ll group of
ncestr
l genes. They w
y they
bind to the lig
nd (the sign
l-rele
sing
molecule)
nd the function
l
nswer of the cell

re specific. When
lig
nd binds to
receptor,

series of re
ctions is initi
ted th
t
lters the
function of the cell. Receptors with direct
nd
indirect lig
nd effects c
n be distinguished. Epinephrine, norepinephrine,
nd h
ist
mine

ct directly
nd very r
pidly. Peptide hormones
such
s insulin or
drenocorticotropic hormone
(ACTH) initi
lly occur
s precursor polypeptides, which
re split by specific prote
ses to
form
ctive molecules. Some peptide hormones

re coded for by
common gene; differenti
l
RNA splicing of the tr
nscript of this gene pro-

duces different precursors for tr
nsl
tion.

(Abbrevi
tions used: ACTH,
drenocorticotropic hormone; FSH, follicle-stimul
ting
hormone; LH, leutinizing hormone; TSH, thyroid-stimul
ting hormone.) (Figure d
t

fter
Lodish et
l., 2000.)
A. Cell surf
ce receptors with direct
lig
nd effect C. Cell surf
ce receptors with indirect
Therefore, if
m
gnitude equ
l to Z is
pplied to e
ch segment of LH
in. such
w
y th
t its centre of gr
vity is
t the centre of the segment,
Elll the m
gnitudes 1 will be equ
l to A. Further, the centre of gr
vity
of the m
gnitude m
de up of
ll these m
gnitudes will he the point E,
remembering th
t they
re
n even numher
nd th
t LE is equ
l to HE
rrClJ}{H;lLIUn V).
Simil
rily it could he shown th
t if
m
gnitude
to Z w
s
pplied to e
ch of the segments of KH, with its centre
.nr.....

t the centre of e
ch segment,
ll those m
gnitudes 1 would
to B
nd th
t the combined centre of gr
vity would he D.
~4']I"'_'I"''I'.I'I.
. A is
pplied
t the point E
nd the m
gnitude B
t
.In.er~~to]re cert
in equ
l m
gnitudes
re pl
ced on the
centres of gr
vity
re sep
r
ted from e
ch other by the

n even number. It is therefore cle
r th
t
m
gnitude composed of
ll these m
gnitudes
..u.:&"'~.""JL'U ~J.L the line on which the centres of gr
vity of
rrO'DO~;ltl()n V). But the length LE is
EC to the length CK. Thus the
lQl~lgIlllttlde composed. of
U these
re
s is the
m
~].tu{le A is
pplied to the . point E
nd
the two
re
s will be in equilibrium
r:!I"I:rlI ... "I:T
e~1te.ld.el
.. thiS J)r~(}D~).slltlOn to. the. c
se
of m
gnitudes
demonstr
tion depended
reproduced this proof of
I
Re
d, "the combin
tion of
ll these m
gnitudes. "28
THE ORIGINS
Proposition VI in its entirety in order to illustr
te the n
ture of Archimedes' logic
l
pp
r
tus. This should not be
llowed, however, to
cre
te too gre
t
n illusion of power.
Indeed, Archimedes
ssumes in this proof th
t the lo
d on the
fulcrum of
lever is equ
l to the sum of the two weights which it
supports.! Further, he m
de use of the principle of superposition
of equilibrium st
tes, without emph
sising th
t this w
s
n experiment
l postul
te. Fin
lly,
nd this is the most telling objection to
the preceeding
n
lysis, Archimedes, together with those of his successors who tried to improve his proof, t
citly m
de the hypothesis
th
t the product PL me
sures the effect of
weight P pl
ced
t

dist
nce L from
horizont
l
xis. In f
ct, in the c
se of complete
symmetry which is envis
ged in Archimedes' first postul
te, equilibrium obt
ins wh
tever l
w of the form Pf(L)is t
ken
s
me
sure
of the effect of the weight P. It is therefore impossible, with the help
of Archimedes' postul
tes
lone, to subst
nti
te Proposition VI in

logic
l w
y.2
For the rest, the tre
tise On the Equilibrium of Pl
nes is concerned
with the determin
tion of the centres of gr
vity of p
rticul
r geometric
l figures. After h
ving obt
ined the centres of gr
vity of
tri
ngle,
p
r
llelogr
m
nd
tr
pezium, Archimedes determined the
centre of gr
vity of
segment of
p
r
bol
by me
ns of
n
n
lysis

which is
milestone in the history of m
them
tics (Book II, Proposition VIII).

We sh
ll now concern ourselves with Archimedes' tre
tise on
Flo
ting Bodies. The
uthor st
rts from the followillg hypothesis" The n
ture of
fluid is such th
t if its p
rts
re equiv
lently
pl
ced
nd continuous with e
ch other, th
t which is the le
st compressed is driven
long by th
t which is the more compressed. E
ch p
rt
;of the fluid is compressed by the fluid which is
bove it in
vertic
l
dir~ction, whether the fluid is f
lling some,vhere or whether it is
driven from one pl
ce to
nother. "
From this st
rting point, the following propositions derive in

logic
l sequence.
Proposition I. - If
surf
ce is intersected by
pl
ne which
lw
ys
p
sses through the s
me point
nd if the section is
circumference
(of
circle) h
ving this fixed point
s its centre., the surf
ce is th
t
of
sphere.
1 This is
point which c
n be est
blished rigorously by consider
tions of symme
try

lone,
s FOURIER w
s to show, much l
ter, in his perfection of
simil
r
ttemp
t due
to n'ALEMBERT.
2 Cf MACH, Mech
nics, p. 21.
Throughout this work, M
ch's tre
tise ,vill be
indic
ted by the letter ]}f.29
HELLENIC SCIENCE
Proposition II. - The surf
ce of
ny fluid
t rest is spheric
l
nd
the centre of this surf
ce is the s
me
s the centre of the E
rth.
This result h
d
lre
dy,
s we h
ve seen, been enunci
ted by Aristotle.
Proposition III. - If
body whose weight is equ
l to th
t of
the s
me volume of
fluid (
) is immersed in th
t fluid, it will sink
until no p
rt of it rem
ins
bove the surf
ce, but will not descend
further.
We sh
ll reproduce the proof of this proposition, which is the
origin of Archimedes' Principle.
" Let
body h
ve the s
me he
viness
s
liquid. If this is possible, suppose th
t the body is pl
ced in the fluid with p
rt of it
bove
the surf
ce. Let the fluid be
t rest. Suppose th
t
pl
ne which
p
sses through the centre of the E
rth intersects the fluid
nd the
K
D
Fig. 3
body immersed in it in such
,y
y th
t the section of the fluid is ABeD

nd the section of the body is EHTF. Let [( be the centre of the
E
rth, BHTC be the p
rt of the body which is immersed in the fluid

nd BEFC the p
rt which projects out of it. Construct
pyr
mid
whose b
se is
p
r
llelogr
m in the surf
ce of the fluid (
)
nd whose

pex is the centre of the E
rth. Let the intersection of the f
ces
of the pyr
mid by the pl
ne cont
ining the
rc ABCD be KL
nd
[(M. In the fluid,
nd below EFTH dr
w
nother spheric
l surf
ce
XOP h
ving the point I(
s its centre, in such
w
y th
t XOP is the
section of the surf
ce by the pl
ne cont
ining the
rc ABCD. T
ke

nother pyr
mid equ
l to the first, with which it is contiguous
nd
continuous,
nd such th
t the sections of its f
ces
re KM
nd KN.
Suppose th
t there is, in the fluid,
nother solid RSQY which is m
de
of the fluid
nd is equ
l
nd simil
r to BHTC, th
t p
rt of the body30
THE ORIGINS
EHTF which is immersed in the fluid. The portions of the fluid
which
re cont
ined by the surf
ce XO in the first pyr
mid
nd the
surf
ce OP in the second pyr
mid
re equ
lly pl
ced
nd continuous
with e
ch other. But they
re not equ
lly compressed. For the

portions of the fluid cont
ined in XO
re compressed by the hody

EHTF
nd
lso by the fluid cont
ined by the surf
ces LM, XO
nd
those of the pyr
mid. The p
rts cont
ined in PO
re compressed
hy the solid RSQ Y
nd by the fluid cont
ined by the surf
ces OP, MN

nd those of the pyr
mid. But the weight of the fluid cont
ined
between MN
nd OP is less th
n the combined he
viness of the fluid
between LM
nd XO
nd the solid. For the solid RSQY is sm
ller
th
n the solid EHTF-RSQY is equ
l to BHTC-
nd it h
s heen

ssumed th
t the hody immersed h
s, volume for volume, the s
me
he
viness
s the fluid. If therefore one t
kes
w
y the p
rts which

re equ
l to e
ch other, the rem
inder will be unequ
l. Consequently
it is cle
r th
t the p
rt of the fluid cont
ined in the surf
ce OP will
be driven
long by the p
rt of the fluid cont
ined in the surf
ce XO,

nd th
t the fluid will not rem
in
t rest. Therefore, no p
rt of the
hody which h
s been immersed will rem
in
bove the surf
ce. However, the body will not f
ll further. For the body h
s the s
me
he
viness
s the fluid
nd the equiv
lently pl
ced p
rts of the fluid
compress it simil
rily."
Proposition IV. - If
body which is lighter th
n
fluid is pl
ced
in this fluid,
p
rt of the 'body will rem
in
bove the surf
ce.
(Proof
n
logous to th
t of Proposition III.)
Proposition V. - If
body which is lighter th
n
fluid is pl
ced
in the fluid, it will he immersed to such
n extent th
t
volume of
:fluid which is equ
l to the volume of the p
rt of the body immersed
h.
s the s
me weight
s the whole body.
The di
gr
m is the s
me
s the preceding one (Proposition III).
" Let the liquid be
t rest
nd the body EHTF be lighter th
n
the fluid. If the fluid is
t rest, p
rts which
re equiv
lently pl
ced
,vill be simil
rly compressed. Then the fluid cont
ined by e
ch of
the <_urf
ces XO
nd OP is compressed by
n equ
l weight. But,
if the body BHTC is excluded, the weight of fluid in the first pyr
mid
is equ
l, with the exclusion of the fluid RSQY, to the ,veight of fluid
in the second pyr
mid. Therefore it is cle
r th
t the weight of the
body EHTF is equ
l to the weight of the fluid RSQY. From which
it follows th
t
volume of fluid equ
l to th
t of the body which is
immersed h
s the s
me weight
s the whole body."HELLENIC SCIENCE
31
Proposition VI. - If
body which is lighter th
n
fluid is tot
lly

nd forcibly immersed in it, the body will be thrust upw
rds with

force equ
l to the difference between its weight
nd th
t of
n equ
l
volume of fluid.
Proposition VII. - If
body is pl
ced in
fluid which is lighter
th
n itself, it will f
ll to the bottom. In the fluid the body will be
lighter by
n
mount which is the weight of the fluid which h
s the
s
me volume
s the body itself.
The first Book of the tre
tise On Flo
ting Bodies concludes with
the following hypothesis- (,(, We suppose th
t bodies which
re thrust
upw
rds
ll follow the vertic
l which p
sses through their centre of
gr
vity. "
In Book II, Archimedes modified the principle which is the subject
of Proposition V, Book I, to the following form" If
ny solid m
gnitude which is lighter th
n
fluid is immersed
in it, the proportion of the weight of the solid to the weight of
n
equ
l volume of fluid will be the s
me
s the proportion of the volume
of th
t p
rt of the solid which is submerged to the volume of the whole
solid. "
We sh
ll p
ss over the proof of this proposition, in which Archimedes
once more deploys th
t powerful logic
l
pp
r
tus with which we
re
now f
mili
r. The rest of Book II is devoted to
det
iled study of
the equilibrium of
flo
ting body which is sh
ped like
right segment

of
(,(, p
r
bolic conoid." In Archimedes' l
ngu
ge (in the tre
tise

On Conoids
nd Spheroids), this term refers to the figure which we
would now c
ll
p
r
bolic cylinder. It m
y be surmised th
t Archimedes w
s interested in this problem for
most pr
ctic
l re
son, for
this surf
ce is simil
r to th
t of the hull of
ship.
It is of interest th
t, throughout this study, Archimedes
pproxim
ted the free surf
ce of
fluid by
pl
ne,
nd th
t he tre
ted vertic
ls

s if they were p
r
llel. This is necess
ry if the concept of centre
of gr
vity is to be utilised. Thus Archimedes must h
ve understood
the necessity
nd the pr
ctic
l import
nce of this
pproxim
tion,
even though his principle w
s b
sed on the convergence of the vertic
ls

t the centre of the E
rth, the spheric
l symmetry of fluid surf
ces

nd
r
ther v
gue hypothesis
bout the pressures obt
ining in the
interior of
fluid.CHAPTER TVVO
ALEXANDRIAN SOURCES
1.
~t\ND
ARABIC MANUSCRIPTS
THE " MECHANICS" OF HERO OF ALEXANDRIA.
It seems th
t Hero of Alex
ndri
lived
t some time during the
lInd Century A. D. His tre
tise Mech
nics discusses cert
in simple
m
chines-the lever, pulley-block
nd the screw-
lone or in v
rious
combin
tions,
nd is only
v
il
ble to us in the form of
n
r
bic version
which h
s been tr
nsl
ted
nd published by C
rr
de V
ux. 1
As f
r
s it concerns the history of mech
nics, the essenti
l import
nce of this work lies in the f
ct th
t its
uthor used the now cl
ssic
l
ide
of moment in his discussion of
lever which w
s not str
ight.
Whether or not this conception w
s
n origin
l one rem
ins doubtful.
Indeed
lex
ndri
n le
rning h
d
ccess to
tre
tise of Archimedes th
t
w
s devoted to levers (lleel evywv)
nd in which the problem of the

ngul
r lever h
d been tre
ted. No tr
ce
of this writing is ext
nt.
However this m
y he, we sh
ll quote
from Hero's Mech
nics.
"Consider
ll
rm of
b
l
nce which
does not h
ve the s
me thickness or
he
viness throughout its length. It m
y
he m
de of
ny m
teri
l. It is in equilibrium when suspended from the point 'Y
-by equilibrium we underst
nd the
rrest
of the be
m in
st
hle position, even
e
though it m
y be inclined in one direction
Fig. 4
or the other. Now let weights he suspended

t some points of the be
m-s
)'"
t ()

nd B. The be
m will t
ke up
new position of equilibrium
fter the
weights h
ve heen hung on. Archimedes h
s shown, in this c
se
s
1
Journ
l
si
tique, 1894.ALEXANDRIAN SOURCES AND ARABIC MANUSCRIPTS
mW]r'se .&.v...
,P,"....
33
"'''''''''''''''.VLL of the weights to e
ch other is the s
me
s the
VLL of the :respective dist
nces. l Concerning these dist
nces
irregul
r
nd inclined he
ms, it should he im
gined th
t

llowed to f
ll from y tow
rds the point ,. Construct

p
sses through the point '-the line 11'O-
nd ""'"hich should

rr
nged to intersect the string
t right
ngles. When the be
m is

t rest the rel
tion of'1J to CO is the s
me
s the rel
tion of the weight

hung
t the point B to the ,veight hung
t the point 6. ."

Hero employed
simil
r
rgument in his discussion of the ,vheel
nd

xle. In f
ct, in reducing the study of these m
chines to the principle
of circles he w
s implicitly using the notion of moment. Thus it is
cle
r, though not explicitly st
ted, th
t in his discussion of the
xle
Hero underst
nds th
t
,veight , c
n be repl
ced by
n equ
l force

pplied t
ngenti
lly
t
bec
use t4F h
s the s
Ine moment
s ~2
Fig. 5
2.
PAPPUS'S THEORIES OF THE INCLINED PLANE AND OF THE CENTRE
OF GRAVITY.
P
ppus (IVth Century A. D.)
ppe
rs to he the only geometer of
Antiquity who took up the problem of the motion
nd equilibrium of

he
vy body on
n inclined pl
ne. The proof th
t we sh
ll
n
lyse
now is t
ken from Book VIII of his Collections (From
mong the v
ried

nd delightful problems of mech
nics) .
P
ppus
ssumes th
t
cert
in effort y is necess
ry to move
weight
t:J.. on the horizont
l pl
ne ltv,
nd th
t
power f) is necess
ry to dr
w it
1 CARRA DE V AUX'S surmise th
t Hero is referring to the tre
tise IIeei 'vywv is
prob
bly correct.
2 Cf. JOUGUET, Lectures de .l\fec
nique, ''''01. I, p. 215. Throughout the prese
nt
book this tre
tise will be indic
ted by the letters L. AI.
:334
THE ORIGINS
IIp the inclined pl
ne /l/~.
He sets out to determine the rel
tion between
y
nd o.
The weight ex. on the pl
ne It'" h
s the form of
sphere with centre e.
P
ppus reduces the investig
tion of the equilibrium of this sphere on
K
Fig. 6
th~
inclined pl
ne to the following problem. A b
l
nce supported
t A
c
tries the weight oc
t e
nd the weight p which is necess
ry to keep
it in equilibrium
t 1J-theend of the horizont
l r
dius ene The l
w of
the
ngul
r lever, which P
ppus borrows from Archimedes' fleel ~vywv
or from Hero's Mech
nics, pro"ides the rel
tion
fJ =
ex.
eC
5:.
rj~
On the horizont
l pl
ne where the power necess
ry to move oc is y,
the power necess
ry to move
long fJ will be
P
ppus then
ssumes th
t the power () th
t is
ble to move the
weight ex. on the inclined pl
ne It'" will be the sum of the powers <5
nd
y, th
t is
() == Y
1ALEXANDRIAN SOURCES AND ARABIC ~iANUSCRIPTS
35
.lvidently the necessity of
power y for pulling the weight ~ on the
~~~~Dt
l pl
ne derives from Aristotle's dyn
mics, in which
ll unn
t~.JDotion requires
motive
gency. The
rgllment by which P
ppus
introduces the lever fAf] supporting the two ,veights (X
nd fJ is r
ther

n
tur
l one, even though it does not le
d to
correct ev
lu
tion of /3..
The l
st hypothesis, concerning the
ddition of b
nd y, the powers
th
t.
re necess
ry to move fJ
nd
respectively on the horizont
l pl
ne
is, on the other h
nd, most str
nge..

However incorrect it m
y h
ve been, this proof ,"-
s destined to

inspire the geometers of the Ren
iss
nce.. Guido Ub
ldo w
s to
dopt
it
nd G
lileo w
s to be occupied in demonstr
ting its f
lsehood .
Archimedes cert
inly formul
ted
precise definition of centre of
gr
vity, but there is no tr
ce of
nything of this kind in those of his
writings th
t
re
v
il
ble to us. Therefore it is of some v
lue to
record the definition which is due to P
ppus.
Im
gine th
t
he
vy body is suspended
hy
n
xis rt./3
nd let it t
ke up its equilibrium position. The vertic
l pl
ne p
ssing through ct/l ,,' will cut the body into
two p
rts th
t
re in equilibrium with e
ch
other
nd which will be hung in such
w
y,
qn one side of the pl
ne
nd on the other,
to be equ
l ~rith respect to their weight. ."
r
king
nother
xis
'p'
nd repe
ting the
Fig. 7
ii
me oper
tion., the second vertic
l pl
ne
p
ssing through
' P' will cert
inly cut the
first-if it were p
r
llel to it " e
ch of these two pl
nes would divide
the body into two equ
l p
rts which would be
t the s
me time of equ
l
weight
nd of unequ
l weight, which is
bsurd.. "
Now suspend the body from
point y
nd dr
w the vertic
l y~
through the point of suspension when equilibrium is est
blished. T
ke

second point of suspension y'
nd, in the s
me w
y, dr
w the vertic
l
,.'3'. The two lines yo, 1"0' necess
rily intersect. For if not, through
e
ch of them
pl
ne could be dr
wn so
s to divide the body into two
p
rts in equilibrium with e
ch other,
nd in such
w
y th
t these two
pl
nes were p
r
llel to e
ch other.. This is impossible.
All lines like yo will therefore intersect
t one unique point of the
body th
t is c
lled the centre of gr
vity.
P
ppus did not distinguish cle
rly,
s Guido Uh
ldo w
s to do in his
Con"ment
r_y on ./lrchimedes' two books on the equilibrium of weights (1588)
between '" equiponder
nt" p
rts, th
t is p
rts th
t
re in equilibrium in
the positions ,\!-hich they occupy,
nd p
rts ,vhieh h
ve the s
me weight.
*836
3.
THE ORIGINS
THE FRAGMENTS ATTRIBUTED TO EUCLID IN ARABIC WRITINGS.
Greek
ntiquity does not
ttrihute
ny work on mech
nics to Euclid.
However his n
me occurs frequently in this connection in the writings
of
r
bic
uthors.
Euclid's book on the b
l
nce,
n
r
bic m
nuscript of 970 A. D.
which h
s been brought to light by Dr. Woepke,l seems to h
ve rem
ined unkno\vn to the western Middle Ages. This relic of greek science m
y
be contempor
neous with Euclid
nd m
y thus
nted
te Archimedes.
It cont
ins
geometric
l proof of the l
w of Ie"vers which is independent
of Aristotle's dyn
mics
nd which m
kes explicit
ppe
l to the hypothesis th
t the effect of
weight P pl
ced
t the end of
n
rm of
lever
is expressed by the product PL. We h
ve h
d occ
sion to emph
sise
the necessity of this hypothesis in our
n
lysis of Archimedes' proof.
The tre
tise Liber Euclidis de gr
vi et levi, often simply c
lled De
ponderoso et levi, h
:5 been known for
long time. It includes
very
precise exposition of
ristoteli
n dyn
mics which is
rr
nged,
fter
Euclid's style, in the form of definitions
nd propositions. The l
tin
text renders the terms ~{)'VCt.Jlq;
nd i
xv;, by which Aristotle me
nt
" power",
s virtus
nd fortitudo. Bodies th
t tr
vel equ
l dist
nces
in the s
me medium-
ir or w
ter-in times which
re equ
l to e
ch
other,
re s
id to be equ
l in t"irtus. Bodies th
t tr
vel equ
l dist
nces
in unequ
l times
re of different virtus,
nd th
t which t
kes the shorter

time is s
id to h
ve the gre
ter virtus. Bodies of the s
me kind
re
those th
t, volume for volume,
re equ
l in virtus. Th
t which h
s
the gre
ter virtus is s
id to be solidius (more dense).
The virtus of hodies of the s
me kind is proportion
l to their dimensions ; th
t is, the bodies f
ll with velocities which
re proportion
l to
their volume. If two he
vy bodies
re joined together, tIle velocity
with which the combin
tion will f
ll will be the sum of the velocities
of the sep
r
te bodies.
Duhem h
s found, in
XIVth Century m
nuscript,2 four propositions on questions in st
tics which complete De ponderoso et levi. This
m
nuscript cont
ins
theory of the rom
n b
l
nce,
nd shows th
t the
f
ct th
t the b
l
nce is
he
vy homogenous cylindel' does not
lter the
rel
tion of the weights to e
ch other.
Fin
lly, in
XIIIth Century m
nuscript, Duhem h
s une
rthed

text c
lled Liber Euclidis de ponderibus secundunt terminorum circonferenti
m 3 which connects the l
w of levers ,vith
ristoteli
n dyn
mics

nd
lso cont
ins
theory of the ronl
n b
l
nce.
1
2
3
Journ
l
si
tique, Vol. 18, 1851, p. 217.
Bibliotheque N
tion
le, P
ris, l
tin collection, 1\ls. 10,260.
Ibid., Ms. 16,649.ALEXANDRIAN SOURCES AND ARABIC MANUSCRIPTS
37
BOOK OF CHARISTION.
Ch
r
stonis is the l
tin version of
n
r
bic text due to
ibn Kurr
h (836-901). The origin
l greek version
:reln
:ms unknown,
nd the question of whether k
r
ston (in Ar
bic II,rsJ~ftn) refers merely to the rom
n b
l
nce or to the n
me of the greek
f!e.)m,etf~r Ch
ristion (
contempor
ry of Philon of Byz
ntium in the
B. C.) h
s been the subject of much schol
rly deb
te.
sh
ll
Duhem 1 in summ
rising the theory of the rom
n
b
l
nce which is found in Liber Ch
r
stonis.
2e(J~me~ter Thitbit
b
d
u
9


e
Fig. 8
A he
vy homogeneous cylindric
l be
m
,b whose
rms
g
nd bg
~:re.
unequ
l m
y he m
int
ined in
horizont
l position by me
ns of

If bd is the
mount
~y which the longer
rm exceeds the shorter
rm
nd u is the centre of
/6
, the weight e ,viII be to bd
s gu is to g
. If p is the tot
l weight of
It the be
m
iw'eight e hung from the end of the shorter
rm
g.
db
e -- p - -,....
--2g

If this weight were known it could be represented ex
ctly by

hung from the shorter
rm,
nd the k
r
ston
rr
nged in this
wfl.ycould be tre
ted
s
weightless he
m.
~. We must
lso mention,
s one of the sources of st
tics, the tre
tise
C
nonio,2
l
tin tr
nsl
tion of
greek text ,vhich
dds nothing
essenti
l to Li ber Ch
r
stonis.

~eAl'C .. p
n
1
2
O. S., Vol. I, p. 90.
Bibliotheque N
tion
le, P
ris, l
tin collection, Ms. 7378 A.CHAPTER THREE
THE Xlllth CENTURY
THE SCHOOL OF JORDANUS
1.
]ORDANUS OF NEMORE AND" GRAVITAS SECUNDUM SITUM. "
The Middle Ages h
d
ccess to the Problems of Mech
nics
nd to the
works of Aristotle. They h
d
lso inherited the fr
gments
ttributed
to Euclid-with the exception of the Book on the B
l
nce-
s well
s
the Liber ClJ,
r
stonis from
r
bic le
rning. They h
d no knowledge
of Archimedes, Hero of Alex
ndri

nd P
ppus.
In spite of the rese
rches of the schol
rs, the person
lity of J ord
nu8
rem
ins mysterious. At le
st three XllIth Century m
nuscripts on
st
tics; h
ve been
ttributed to him,
lthough these
re cle
rly in the
style qf different
uthors. Neither Jord
nus's n
tion
lity nor the period id which he lived is known with
ny cert
inty. D
unou believes
him t9 h
ve lived in Germ
ny
bout 1050, Ch
sles
ssoci
tes him with
the ~IIIth Century while Curtze pl
ces him
bout 1220 under the n
me
of Jbrd
nus S
xo. Mich
ud h
s identified him with R
imond Jord
n,
provost of the church of Uzes in 1381 which is cle
rly too l
te. With
Montucl
, we sh
ll here
dopt the intermedi
te vie,v th
t
ssoci
tes
Jord
nus of Nemore with the XIIlth Century.
Like Duhem, we sh
ll follow the Element
J ord
ni super demonstr
tionem ponderis. 1 This work comprised seven
xioms or definitions
foll().wed by nine propositions. The essenti
l origin
lity of Jord
nus l
y
in the system
tic use, in his study of the motion of he
vy bodies, of the
effective p
th in
vertic
l direction
s
me
sure of the effect of
weight,
which w
-s usu
lly pl
ced
t the end of
lever
nd described
circle in
consequence. Thus his st
tics stems, implicitly, from the principle of
virtu
l work. The word work, t
ken in the modern sense., is to be con1 Bibliotheque N
tion
le, P
ris, Ms. 10,252, d
ted 14.64.
There
lso exists
n in
complete m
nuscript of the s
me work, d
ting from the XIlIth Century, in the Bib
lio
theque M
z
rine, Ms. 3642.39
THE XI11th CENTITRY
tr
sted with the word velocity
nd with the concept of virtu
l velocities
l\rhich m
y he tr
ced in the
rguments of Problems of Mech
nics. Of
course lord
nus never used the word" work" itself.. He considered
the he
viness of
p
rticle rel
tive to its situ
tion (gr
vit
s secundum
situm) without m
king cle
r the rel
tion th
t exists between this qu
ntity
nd the he
viness in the strict sense.
Jord
nus formul
ted his principle in
picturesque L
tin which merits
quot
tion.
" Omnis ponderosi mot
m esse
d lnedium, virtutemque ipsius potenti
m
d inferior
tendendi et motui contr
rio resistendi.
" Gr
vius esse in descendendo qu
ndo ejusdem motus
d medium rectior.
" Secundum situm gr
vius, qu
ndo in eodem situ minus obliquus est
descensus.
" Obliquiorem
utem descensum in e
dem qu
ntit
te minus c
pere de
directo. "
Or" The motion of
ll he
vy things is tow
rds the centre,! its strength
heing the power ,vhich it h
s of tending do,vnw
rds
nd of resisting

contr
ry motion.
,,4 A moving body is the he
vier in its descent
s its motion tow
rds
the centre is the more direct.

'" A body is the he
vier bec
use of its situ
tion
s, in th
t situ
tion,
descent is the less oblique.
,~ A more oblique descent is one th
t, for the s
me p
th, t
kes less
of the direct. "
Thus
cert
in weight pl
ced
t b,

t the end of the lever cb, h
s
sm
ller
gr
Vity secundum situm th
n the s
me
~~~ght h
s when it is
t
,
t the end
of the horizont
l r
dius c
. Indeed, on

....- - - -.. c
the circumference of the circle with
centre c
nd r
dius c
== cb, if the
- - --~z'
:b,ody f
lls from b to It
long the
rc
I
bii the effective p
th in
vertic
l
I
direction is b' h'. On the other h
nd
if. ,the body st
rts from

nd f
lls
Fig. 9

long
n ~
rc liZ., ","hich is equ
l to
the
rc bh, the effective vertic
l p
th
is cz'
nd is gre
ter th
n b' h'. Thus the descent bh, equ
l to the
descent liZ., is more oblique th
n th
t
nd t
kes less of the direct.
,
1
ITnderstood
s the common centre of
ll he
vy things in . ' . \'ristotle's sense
.40
THE ORIGINS
This ide
led J ord
nus to
proof of the rule of the equilibrium of the
str
ight lever whose origin
lity c
nnot be contested.
d
b
" Let
cb he the he
rn,

nd b the weights th
t it c
rries,
nd suppose
th
t the rel
tion of b to
is the s
me
s th
t of c
to cb. I m
int
in
th
t this rule will not ch
nge its pl
ce. Indeed, if the
rm supporting b
f
lls
nd the he
m t
kes up the position dee, the weight b will descend
by he ~nd
will rise by fd. If
weight equ
l to the weight b is pl
ced

t 1,
t
dist
nce such th
t cl == eb, this will rise in the motion by
gm === he. But it is cle
r th
t dfis to mg
s the weight I is to the weight
.
Consequently, wh
t is sufficient to bring
to d will be sufficient to bring
I to m. But we h
ve shown th
t b
nd 1 counterb
l
nce e
ch other ex
ct~y, so th
t the supposed motion is impossible. This will
lso he true
jof the inverse motion. "
Duhem writes in this connection 1 _
'" Underlying this demonstr
tion of Jord
nus the following principle
is cle
rly evident-th
t which c
n lift
weight to
cert
in height c
n

lso lift
weight which is k times
s
8
c
gre
t to
height which is k times less.
This principle is then the s
me
s th
t
which Desc
rtes took
s
b
sis for his
complete theory of st
tics
nd which,
th
nks to John Bernoulli, bec
me the
principle of virtu
l work. "
Jord
nus w
s less fortun
te "\\Then he
turned his
ttention to the
ngul
r lever.
He considered
lever
c! c
rrying equ
l
f

weights
t

nd f which were pl
ced in

Fig. 11
such positions th
t
c == ef. Jord
nus
w
s of the opinion th
t, under these
1
o.
S., Vol. I, p. 121.THE Xlllth CENTURY
41
would domin
te f. He
rrived
t this conclusion by conequ
l
rcs ;;1
nd ;f. It is cle
r th
t the "Co direct " t
ken

is gre
ter th
n the " direct " t
ken by the weight f.
conclusion is obt
ined hec
use, since the link
ges
re
two displ
cements;;]
nd j;;"
rc not simult
neously possible.
thus misunderstood the ide
of moment.

s the XIIIth Century the Element
Jord
ni were gener
lly
copyi:;ts with lJe C
nonio to form the Liber Euclidis de
This
rtifici
l
ssoci
tions
nd this im
gin
tive titles
desp
ir of the schol
rs
nd it h
s needed
ll the le
rning of
................,.,. . . . . to elucid
te them.
Every truly novel ide
evokes
re
ction. The Element
Jord
ni
did not provide
n exception to this rule. In the XIIIth Century

eritie wrote
comment
ry of J ord
nus which Duhem c
lls the Perip
tetic Con~n~ent
ry.2 This
uthor
t every turn invokes the
uthority of
Aristotle
nd h
s scruples
hout
pplying the gr
vit
s secundum situm
to .8 motionless point-in modern l
ngu
ge,
bout m
king
ppe
l to

mtu
l displ
cement. It does not
ppe
se his conscience to consider
th
t rest is the end of motion. "" The scientific v
lue of the Comment
ry is nothing," decl
res Duhem. 3 "But its influence did not
dis
ppe
r for
very long time,
nd even the gre
t geometers T
rt
gJi
, Guido Uh
ldo
nd Mersenne h
d not entirely freed themselves
ftom it. "
2}~ THE ANONYMOUS AUTHOR OF "LIBER JORDANI DE RATIONE PON..
,
DERIS .. "
TIlE ANGULAR LEVER.
THE INCLINED Pl,ANE.
We now come to
n especi
lly noteworthy work which figures in the
s
me m
nuscript
s the Perip
tetic Comment
ry under the title Liber
Jii,i!
,ni de r
tione ponderis,
nd which did not rem
in unknown in the
Ren
iss
nce. l'
rt
gli
sent it to Curtius Troj
nus who puhlished it
in 1565. This work supercedes
nd rectifies the Element
Jord
ni on
m~ny import
nt points. ...J\ll the s
me, it is h
sed on the s
me principle
of gr
vit
s secundum situm.
Duhem, ,\Tho brought this m
nuscript to light, terms the
nonymous

uthor
H Precursor of Leon
rdo d
Vinci." Indeed, in m
ny respects
this "precursor surp
ssed Leon
rdo, who, for ex
mple, spent himself
in fruitless efforts to ev
lu
te the
pp
rent weight of
hody on
n
inclined pl
ne.. It seems more n
tur
l to simply spe
k of
n
nonymous
1
2
3
Bibliotheque N
tion
le, P
ris, l
tin collection., Mss. 7310
nd 10.,260.
Ibid., 1\'15. 7378 A.
O. S.') 'Tol. I, p. ]34.42
THE ORIGINS

uthor of the XIIIth Century,
disciple of Jord
nus who h
d out..
stripped his m
ster.
In connection with the hent lever this
uthor corrected Jord
nus's
error. As before, let
lever
cf c
rry equ
l ,veights
t

nd f
nd
be pl
ced in such
position th
t

' == if'.

c
I
It is impossible th
t the weight
should domin
te the weight f.
!For if two
rcs ;;}", fl,
re considered on the two circles dr
wn through


nd f
nd corresponding to equ
l
ngles -;;J"
nd fii; the descent
of

long rh necessit
tes th
t the equ
l weight
t f should rise through

dist
nce In which is gre
ter th
n rh. This is impossible.
In the s
me \v
y it c
n be seen th
t f will not domin
te
. For
if the
rcs j;
nd ~ correspond to equ
l
c

nglesft;
nd;;;;;;' the descent ofj
long
tx m
kes it necess
ry th
t the equ
l
I
weight pl
ced
t
should rise by pm,


~------~ J


\vhich is gre
ter th
n tx. This is imposI
sible. Therefore there is equilibrium in
I
the position considered, in which

' if'.
J
The
nonymous
uthor gener
lised
b l - b
this result to
n
ngul
r b
l
nce
I
Fig. 13
c
rrying unequ
l weights
t

nd b,43
THE XIIIth CENTURY

nd obt
ined the result th
t in equilibrium it is necess
ry th
t the
dist
nces

'
nd bb' from

nd b to the vertic
l dr
wn through the
point of support, c,
re in inverse r
tio to the weights

nd b..
We see th
t this
uthor knew
nd used the notion of moment..
Elsewhere he wrote on this subject., '" If
lo
d is lifted
nd the
length of its 81lpport is known, it c
n be determined how much
this lo
d weighs in
ll positions. The weight of the lo
d c
rried
t
e by the support be will he to the weight c
rried
t f by fb
s el is
to fr or
s pb is to xb. .A. ,veight pl
ced
t e,
t the end of the
lever be, will weigh
s if it were
t
u on the lever bj: "
Thus the ide
of gr
vit
s secundum
situm, which Jord
nu8 h
d used qu
lit
tively, bec
me precise.
Our
nonymous
uthor
lso concerned himself with the st
bility of
the h
l
nce,
nd rectified cert
in
errors which were cont
ined in the
relev
nt p
rts of Problems of Mech
nics.
C .....
p - -.. b
More th
n this, he resolved the
Fig. 14
problem of the equilibrium of
he
vy
h~dy on
n inclined pl
ne,
problem
wbich h
d eluded the wisdom of the greek
nd
lex
ndri
n geometers..
, In order th
t this m
y be done, it is first observed th
t the gr
vit
s
stcull,dum situm of
weight on
n inclined pl
ne is independent of
its position on the pl
ne. The
uthor then
ttempts
comp
rison of

, the v
lue th
t th
t gr
vity t
kes on differently inclined pl
nes. We

sh
ll quote from Duhem's tr
nsl
tion of this s
me Xlllth Century
m
nuscript.
'" If two ,veights descend by differently inclined p
ths,
nd if they

re directly proportion
l to the declin
tions, they ,,,ill be of the s
nle
strength in their descent..
,<. Let
b be
horizont
l
nd bd,
vertic
l.
Suppose th
t two
oblique lines d

nd de f
llon one side
nd on the other of bd,
nd
th
t de h
s the gre
ter rel
tive ohliquity.. By the rel
tion of the ohliquities I underst
nd the rel
tion of the declin
tions
nd not the rel
tion
of the
ngles; this me
ns the rel
tion of the lengths of the n
med
lines counted
s f
r
s their intersection with the horizont
l, in such

w
y th
t they t
ke simii
rily of the direct.
" In the second pl
ce, let e
nd h be the weights pl
ced on dc
nd
d
respectively,
Dd suppose th
t the ",-eight e is to the ,veight h
s
,
-...r-----.. .44
THE ORIGINS
d
-----,.,j
...----......---------..
k ....
C
Fig. 15
de is to d
. I m
int
in th
t in such
situ
tion the two weights will
h
ve th.e s
me strength.
" Indeed, let dk be
line h
ving the s
me obliquity
s de
nd,
on th
t line, let there be
weight g which is equ
l to e.
" Suppose th
t the weight e should descend to I, if th
t is possible.,

nd th
t it should dr
w the weight h to m. (It is cle
r th
t the
uthor
im
gines the two weights to he connected by
thre
d which p
sses
over
pulley
t d.) T
ke gn equ
l to hm,
nd consequently equ
l
to el. Dr
w
perpendicul
r to db which p
sses through g
nd h,
s
y ghy. Drop
perpendicul
r It from the point I onto db. Then
drop [the perpendicul
rs] nr, mx,
nd ez. The rel
tion of nr to ng is
th
~ of dy to dg
nd
lso th
t of db to dk. Therefore mx is to nr
s
dk i$ to d
; th
t is to s
y,
s the weight g is to the weight h. But
s e
is nJ)t
ble to pull g up to n (nr ::=: ez), it is no better
hle to pull h up
to /m. The weights therefore rem
in in equilibrium. "
This demonstr
tion, which le
ds to the correct l
w of the
pp
rent
he
viness of
body on
n inclined pl
ne, w
s directly inspired hy th
t
"of Jord
nus concerning the equilibrium of
str
ight lever. Like
th
t, it, implicitly proceeds
ccording to the principle of virtu
l work.
We sh
ll now give some indic
tion of the ide
s on dyn
mics which
were used by the
uthor of Liber Jord
ni de r
tione ponderis.
The environment's resist
nce to the motion of
hody depends
on the sh
pe of the body, which penetr
tes the environment the hetter

s its sh
pe is the more
cute
nd its figure the more smooth. It
depen.ds~ in the second pl
ce, on the density of the fluid tr
versed. All
medi

re compressihle; the lower str
t
, compressed by the upper
ones.,
re the denser
nd those which hinder motions more. At the
front of the moving body will be
p
rt of the medium cQmpressed
on,
nd sticking to it. But the other p
rts of the medium, which

re displ
ced by the mo
lig
nd effect
M
ny hormones c
nnot p
ss through the
pl
sm
membr
ne; inste
d, they inter
ct with
C
CONTABI LITATEA FORM

 DE BAZ
A EVIDE
EN EI EC ONOMICE
E
1.1
1. Obiective e
L

sfritul
c cestei unit i de nv
r re studenii vor fi c
p
b bili s:
define e
sc no iun
ne
de cont

bilit
te;
n ele

g necesi it
te
org
n nizrii con nt
bilit ii l
nivelul fiecrei
unit i p
trimoni

le;
n ele

g rolul pe c
re l
re cont

bilit
te
n n c
drul
ctivit ii


desf ur
te de un
nit ile p
tri imoni
le;

n ele

g modul de
d org
niz
r re
cont
bil lit ii l
nive el microeco onomic.
1.2. Definir re
, necesit t
te
i rolu
ul cont
bili it ii
C
Cont
bilit
t
te

exist

t din cele e m
i vech hi timpuri. C.G. Dum
mitrescu, n Istori

cont
bil lit ii,
rt

c grecii
u

mprumu ut
t tehnic
eviden ei co ont
bile de l
egipteni, i
r de l

ei
u pr relu
t-o rom
m
nii. D
r se p
re c
eviden ele cont
bile sunt
s
mult m
i
m vechi n n istori

omeniri ii. Cont
bil lit
te
n p

rtid dubl s-
nscu ut c
urm

re
pr
cti cii c
ont
bil lilor din
Vene i

i Genov

. n
nul 1 494, Luc
P
ciolo des scrie cont
b
bilit
te
n p
rtid
p
dub bl ntr-o
lucr
re de m
tem

tic i geom
metrie. Dup
p
p
ri i

cestei luc crri,
plic
r re
cont
bi ilit ii n
p
rtid dubl s-
r spndit i n

lte ri
le Europei.
A
Astzi,
con
nt
bilitii i i revin s
rci ini din ce n n ce m
i gr rele. E
c
u
ut s-i dep pe
sc
limitele , fiind pus n situ
i
de
descrie e org
niz
i ii din ce n ce
c m
i comp
plexe c
re opere
z
o
ntr-un m
mediu econ nomic i soc ci
l n contin
nu mic
re e i tr
nsfor rm
re.
Princip
p
lele
spec cte
c
ins strument de descriere, de
d model
re e
ntreprin nderilor;
sub c
r re trebuie
c
ins strument de prelucr
re
inform
ii ilor;
studi
t cont
bilit t
te
c
pr r
ctic s
u u joc soci i
l, nscri s ntr-o r ree
de

(dup N.
N Fele
g) )
restric cii regleme ent
re m
i mult
m s
u m
i puin stric cte.
10Cont
bilit
te
po
te fi consider
t drept o
rt, o tehnic s
u o tiin, d
r indiferent
de
cum
m privi-o, cont
bilit
te
este un joc soci
l ce
re drept fin
lit
te reprezen
t
re
unei
re
liti c
re este entit
te
.
Cont
bilit
te
studi
z
cele l
turi
le reproduciei soci
le c
re se pot exprim
n
et
lon bnesc. E
urmrete existen
i din
mic
p
trimoniului
genilor economici,
procesele economice, pe c
re
ceti
le org
nize
z, st
bilind i nregistrnd rezult
tele
fin
nci
re fin
le.
1.3. Org
niz
re
evidenei cont
bile l
nivel microeconomic
Conform Legii Cont
bilitii nr. 82/1991, ntreprinderile
u oblig
i
s org
nizeze i
s conduc cont
bilit
te proprie, n limb
romn i n moned
n
ion
l. Org
niz
re

cont
bilitii reprezint deci, nu num
i o necesit
te,

cum
m
rt
t
nterior, d
r i o
oblig
ie impus prin reglementrile leg
le n vigo
re.
De
ltfel, entit
te
c
sistem complex economico-soci
l i
dministr
tiv-org
niz
t
oric,
ndeplinete o serie de funcii, n c
drul cror
un rol eseni
l l
re funci
fin
nci
rcont
bil.
Pentru ndeplinire

cestei funcii i n
cel
i timp pentru respect
re
legii, n c
drul
entit ii se org
nize
z i funcione
z un comp
rtiment speci
liz
t, fin
nci
r-cont
bil. n
c
drul
cestui comp
rtiment lucre
z perso
ne cu studii de speci
lit
te (medii i su
perio
re),

vnd
tribuii distincte n domeniul evidenei cont
bile oper
tive i gener
le.
Comp
rtimentul fin
nci
r-cont
bil se subordone
z cont
bilului ef, c
re
re studii
superio
re
n fin
ne-cont
bilit
te.
Aici se consemne
z zilnic i lun
r, to
te oper
iunile economico-fin
nci
re ce
u loc
n entit
te, respectiv: cumprri, vnzri, consumuri, s
l
rii, nc
sri, pli etc., conduc
fin
l l
determin
re
rezult
tului
ctivitii i l
ntocmire
situ
iilor fin
nci
re
nu

le de
sintez i r
port
re cont
bil.
Lun
c
lend
ristic po
rt denumire
de perio
d de gestiune, i
r
nul c
lend
ristic,
de exerciiu fin
nci
r.
n b
z
d
telor furniz
te de eviden
cont
bil, se pot efectu

n
lize economicofin
nci
re privind corel
re
resurselor
loc
te cu rezult
tele obinute, se pot c

lcul
diveri
indic
tori i se po
te determin
evolui
diverselor fenomene n timp, cu f
ctorii poz
itivi i
neg
tivi c
re le-
u gener
t.
11n c
drul entit ii, cont
bilit
te
se org
nize
z pe dou circuite p
r
lele: cont
bilit

te
fin
nci
r i cont
bilit
te de gestiune.
este reglement
t prin norme unit
re;
ofer o viziune glob
l
supr

ctivitii;

re un obiectiv fin
nci
r reflect
re
im
ginii fidele

p
trimoniului;
Cont
bilit
te
fin
nci
r
genere
z fluxuri de inform
ii i documente externe;

plic reguli norm
tive;
ofer d
te utiliz
torilor externi (furnizori, clieni, bnci,
investitori, org
ne de control etc.);
se refer l
perio
de nchei
te (lun,
n).
se l
s l
l
titudine
fiecrei entit i;
ofer o viziune det
li
t
supr

ctivitii;

re un obiectiv economic constnd n supr
veghere
i
controlul
ctivitii prin intermediul costurilor;
Cont
bilit
te
genere
z fluxuri de inform
ii interne;

de gestiune
plic reguli st
bilite n c
drul entit ii;

ofer d
te conducerii entit ii;
se refer l
prezent i viitor (previziuni);
cl
sific cheltuielile dup locul de re
liz
re i destin
i

lor.
12Dup p
rcurgere

cestei uniti de nv
re trebuie s reinei:
Ce este cont
bilit
te
i c
re este rolul
cestei
n c
drul unei
entiti.
C
re sunt circuitele de org
niz
re
le cont
bilitii n c
drul
unei entiti.
Ce este cont
bilit
te
fin
nci
r i prin ce se c
r
cterize
z.
Ce este cont
bilit
te
fin
nci
r i prin ce se c
r
cterize
z.
13UNITATE
EA DE NV
V ARE 2
OBIEC
CTUL I METODA
M
C
CONTABIL
LIT II
2.1
1. Obiective e
L

sfritul
c cestei unit i de nv
r re studenii vor fi c
p
b bili s:
identi fice obiectu ul de studiu
l cont
bili it ii;
identi fice proced deele ce
p
r r in metode i cont
bilit ii;
identi fice i s n n ele
g rolu ul principiil or cont
bili it ii;
identi fice func ii ile cont
bil lit ii i ne ecesit
te

p
plicrii
ce estor
n
cont
b bilit
te
fin
nci
r i n n cont
bilit

te
de gest tiune.
2 Obiect tul cont
bil lit ii
2.2.
C orice di sciplin tii in ific, con nt
bilit
te
reprezint simult
n
C

s
o teorie
t
i me etod. n
c
lit
te

s
de teor rie tiin ific c, cont
bil it
te
repre ezint un sis stem de pr
i incipii i cu unotin e
c
re ex xplic i inf forme
z, i
r
i c
meto
od s
u teh hnic, un
n ns
mblu co oerent de procedee,
p
instrume ente prin c

re se observ v i nregis stre
z resu ursele econo omice
le so
ociet ii, sep p
r
te c

utilit i p
trimoni
l le.
Concep
p ii cu
1. Concep p i

dmin nistr
tiv consider c obiectu ul cont
bil lit ii l
privire l
constit tuie reflect
re
i con ntrolul, n expresie v
loric,
v

f
ptelor
delimit

re

dmini istr
tive, n n vedere
sp prijinirii m

n
gementu ului, pen
tru
ob ine
i defin
nire
cu un minim de eforturi (ch heltuieli) un n m
xim de e efecte eco onomice
obiectu

ului (rezult t
te-venituri i).

cont
bi ilit ii
2. Concep p i
juridic c consider r c obiec ctul cont
b bilit ii l forme
z
f
p
trim
moniul unui i subiect de
d drept (

gent econo
omic), prin n prism

rel
iil or juridice e,
dic
drepturil lor i obl lig
iilor p pecuni
re
(m
ter ri
le)
le unei
u
perso

ne fizice s
u juridic ce, n core el
ie cu
obiecte ele,
dic cu
u bunurile i
v
lorile co orespunzto o
re.
3. Concep p i

econ omic
s
u u
fin
nci


r
consid
der
c
obiectul
cont
b bilit ii l co
onstituie cir rcuitul c
pit t
lului, priv vit
tt sub
spectul
destin

iei lui, c t i sub for rm
modulu ui de dobn ndire, respe ectiv sub
form
de c
pit
l propriu
p
i c

pit
l strin. .
16 Obiectul cont
bilit ii este reprezent
t de reflect
re
n expresie bne
sc

bunurilor mobile i imobile, inclusiv solul, bog iile n
tur
le,
zcmintele i
lte bunuri cu poten i
l economic, disponibilit ile bneti,
Concluzie
titlurile de v
lo
re, drepturile i oblig
iile
gen ilor economici, precum
i micrile i modificrile intervenite n urm
oper
iunilor p
trimoni
le
efectu
te, cheltuielile, veniturile i rezult
tele ob inute de
ceti
.
Obiectivul fund
ment
l
l cont
bilit ii l reprezint furniz
re
de
inform
ii utile lurii deciziilor menite s
sigure o im
gine fidel

p
trimoniului, situ
iei fin
nci
re i
rezult
telor ob inute.
Din
ce
st defini ie rezult c p
trimoniul presupune existen

dou condi ii de
b
z: un titul
r de p
trimoniu (subiect de drept) pe de o p
rte, i
r pe de
lt p
rt
e bunurile i
v
lorile economice, privite c
obiecte de drepturi i oblig
ii, precum i titul
rul d
e p
trimoniu.
Structur
de
ns
mblu
p
trimoniului este urmto
re
:
PATRIMONIU
BUNURI ECONOMICE
PATRIMONIU PROPRIU
(DREPTURI)
PATRIMONIU STRIN
(OBLIGA II)
Bunurile economice se concretize
z n bunuri m
teri
le (terenuri, cldiri, util
je,
mijlo
ce de tr
nsport, stocuri de m
terii prime etc.) i nem
teri
le (proiecte, pr
ogr
me
inform
tice etc.) formnd
vere
entit ii.
Rel
iile de drepturi se refer l
f
ptul c entit
te
i procur o p
rte din
vere din
surse proprii, i
r bunurile procur
te i
p
rin de drept.
Rel
iile de oblig
ii se refer l
f
ptul c entit
te
i procur o p
rte din
vere din
surse mprumut
te, fiind oblig
t s restituie terilor contr
v
lo
re
bunurilor procur

te.
Cont
bilit
te
se ocup cu reflect
re
n expresie v
loric
p
trimoniului, e

nregistre
z circuitul elementelor p
trimoni
le n condiii concrete de timp i sp
iu,
c
lcule
z mrime

cestor elemente i reflect mic
re
p
trimoniului prin oper
iuni de
intrri i ieiri 1 .
1
Ghe. T
l
ghir, Ghe. Negoescu Cont
bilit
te
pe nelesul tuturor. Editur
All, Bucur
eti, 1998.
17Cont
bilit
te
studi
z modul de gestion
re
p
trimoniului, fund
mente
z
deciziile referito
re l
fin
n
re
i utiliz
re
elementelor p
trimoni
le, controle

z re
liz
re

deciziilor i st
bilete rspunderi privind integrit
te
i dezvolt
re
p
trimoniului.
De
semene
, cont
bilit
te
studi
z echilibrul glob
l
l p
trimoniului, prin
respect
re
ecu
iei p
trimoni
le:
BUNURI ECONOMICE = DREPTURI + OBLIGA II
cu deriv
tele s
le:
Drepturi = Bunuri economice Oblig
ii
i
Oblig
ii = Bunuri economice Drepturi
n consecin, entit
te
reprezint o unit
te p
trimoni
l,
l crei p
trimoniu po
te fi
privit sub dublu
spect:
l mijlo
celor economice (bunurile/
vere
) i
l surselor
de
procur
re

cestor mijlo
ce (c
pit
lul) proprii i strine.
Mijlo
cele economice definesc
ctivul p
trimoni
l, i
r sursele definesc p
sivul
p
trimoni
l.
n derul
re
oper
iunilor economico-fin
nci
re,
p
r i o serie de procese
economice, sub form
veniturilor i cheltuielilor, c
re
jut l
nregistr
re
creterii
s
u
diminurii p
trimoniului.
n
cest context, ecu
i
p
trimoni
l de m
i sus, devine:
AVERE = CAPITAL
182.3. Metod
cont
bilit ii
D
torit complexit ii obiectului de studiu, metod
cont
bilit ii reunete m
i multe
procedee tehnice de lucru.
Un
ns
mblu de procedee
fl
te ntr-o strns corel
ie i
Metod
intercondi ion
re c
un tot unit
r, n vedere
st
bilirii normelor i
cont
bilit ii principiilor cu c
r
cter speci
l pe c
re se fund
mente
z cont
bilit
te


i cu
jutorul cror
cercete
z st
re
i mic
re
elementelor
p
trimoni
le
le unit ilor p
trimoni
le.
O tr
stur c
r
cteristic
metodei cont
bilit ii este
cee

folosirii unor procedee
c
re s permit nregistr
re
numeric, cifric,
existen ei i micrii p
trimoniului unit
economice i soci
le n expresie v
loric. Gener
liznd, se po
te spune c metod

cont
bilit ii reprezint tot
lit
te
procedeelor interdependente, pe c
re le folosete

ce
st
n
scopul cuno
terii situ
iei p
trimoniului i
rezult
telor ob inute.
1. procedee comune tuturor tiin elor;
Procedee utiliz
te 2. procedee specifice metodei cont
bilit ii;
de cont
bilit
te 3. procedee
le metodei cont
bilit ii, comune i
ltor discipline
economice.
Procedee comune Observ
i
este f
z
ini i
l
cercetrii obiectului de studiu
l
tuturor tiin elor oricrei tiin e i este utiliz
t pentru cuno
tere
oper
iilor economic
c
re se pot exprim
v
loric i pe c
re le reflect cifric, numeric, cu

jutorul procedeelor s
le specifice.
R
ion
mentul se
plic de metod
cont
bilit ii, pentru c pe b
z
de judec i logice, pornind de l
fenomenele i procesele economice
c
re intr n obiectul su de studiu, s
jung l
concluzii noi (ex.:

ctivul este eg
l cu p
sivul, pentru c ntre mijlo
cele economice i
sursele de fin
n
re

cestor
exist o eg
lit
te perfect).
Comp
r
i
se folosete de metod
cont
bilit ii prin
ltur
re

dou s
u m
i multe fenomene i procese economice c
re se pot exprim

19v
loric, cu scopul de
st
bili
semnrile i deosebirile dintre ele, c


stfel s se tr
g o serie de concluzii. Se folosete frecvent pentru
se
comp
r
veniturile i cheltuielile pe b
z
cror
se st
bilesc rezult
tele
fin
le.
Cl
sific
re

c iune
de mpr ire, distribuire, rep
rtiz
re sistem
tic
pe cl
se s
u ntr-o
numit ordine
obiectelor n func ie de
semnrile
i deosebirile dintre ele. Asemnrile le
propie i le nc
dre
z n

cee
i cl
s, i
r deosebirile le diferen i
z i le distribuie n cl
se
diferite.
An
liz
procedeu tiin ific de cercet
re
unui ntreg,
unui fenomen
c
re se b
ze
z pe ex
min
re
fiecrui element component n p
rte.
Sintez
c
procedeu tiin ific de cercet
re
fenomenelor se b
ze
z
pe trecere
de l
p
rticul
r l
gener
l, de l
simplu l
compus pentru

se
junge l
gener
liz
re.
Procedee specifice Bil
n ul st l
b
z
dublei reprezentri
p
trimoniului n
metodei cont
bilit
te. El furnize
z inform
ii gener
le privito
re l
situ
i

cont
bilit ii economic i fin
nci
r
unei entit i, d
r reflect i l
rel
iile ei
economice cu
lte entit i, fiind complet
t de o serie de situ
ii
nex
prin c
re se explic i se det
li
z
numite l
turi
le
ctivit ii
economico-fin
nci
re
le societ ii.
Contul se deschide n cont
bilit
te
curent pentru reflect
re

fiecrui element din p
trimoniu. Cont
bilit
te
dispune de un sistem
de conturi n c
re reflect
re
oper
iilor economice rezult
te din
mic
re
elementelor p
trimoni
le
re l
b
z dubl
nregistr
re.
B
l
n
de verific
re
sigur n cont
bilit
te
dublei reprezentri i

dublei nregistrri, g
r
n i
ex
ctit ii nregistrrilor efectu
te n
conturi. D
tele b
l
n ei de verific
re st
u l
b
z
ntocmirii bil
n ului.
B
l
n
de verific
re ndeplinete
tt o func ie de control, ct i o
func ie economic, constituind punte
de legtur ntre cont i bil
n .
20Procedeele metodei Document
i
orice oper
ie economic i fin
nci
r referito
re l

cont
bilit ii comune existen
i mic
re
elementelor p
trimoni
le trebuie s fie
i
ltor discipline consemn
t n documente c
re f
c dov
d
nfptuirii lor.
economice Ev
lu
re
procedeul prin c
re d
tele cont
bilit ii sunt reprezent
te
printr-o singur unit
te de msur, crend posibilit
te
centr
lizrii
lor cu
jutorul b
l
n elor de verific
re i gener
liz
re
cu
jutorul
bil
n ului. Ev
lu
re
const n tr
nsform
re
unit ilor n
tur
le n
unit i monet
re cu
jutorul pre urilor.
C
lcul
i
este strns leg
t de ev
lu
re c
procedeu
l metodei
cont
bilit ii. Acest procedeu i gsete
plic
re
ce
m
i l
rg n
domeniul c
lcul
iei costurilor de produc ie.
Invent
riere
se folosete pentru
se cuno
te situ
i
re
l

p
trimoniului reflect
t n cont
bilit
te, i trebuie s se verifice
existen
f
ptic
tuturor elementelor s
le, n scopul descoperirii
neconcord
n elor dintre d
tele nregistr
te n conturi i re
lit
te
de
pe teren.
2.4. Principiile cont
bilit ii
Pentru
se oferi o im
gine fidel
p
trimoniului,
situ
iei fin
nci
re i
rezult

telor
ob inute de ctre entit
te, trebuie respect
te cu bun credin regulile privind ev
lu
re

p
trimoniului i celel
lte norme i principii cont
bile.
Principiul
nu sunt
dmise supr
ev
lu
re
elementelor de
ctiv i

pruden ei veniturilor, respectiv subev
lu
re
elementelor de p
siv i

cheltuielilor, innd cont de deprecierile, riscurile i pierderile
posibile gener
te de desfur
re

ctivit ii unit ii;
pruden
presupune
nticip
re
efectelor unor
c iuni i n speci
l

tr
nsferului de propriet
te cu efecte posibile
supr
exerci iului
s
u
celor p
rcurse dej
, ntruct
supr
lor nu se m
i po
te
interveni din punct de vedere cont
bil;
l
ncheiere
fiecrui exerci iu fin
nci
r se cont
bilize
z

d
toriile i pierderile prob
bile.

21Principiul
continuit
te

plicrii regulilor i normelor privind ev
lu
re
,
perm
nen ei nregistr
re

n
cont
bilit
te
i
prezent
re

elementelor
metodelor p
trimoni
le i
rezult
telor
sigurnd comp
r
bilit
te
n timp

inform
iilor cont
bile;

sigur
plic
re
pentru
cele
i elemente, stucturi, domenii de

ctivit
te,

celor
i metode de l
un exerci iu l

ltul, excluznd
schimb
re
metodelor n cursul exerci iului;
modific
re
metodelor de l
un
n l

ltul trebuie s fie
determin
t de o profund motiv
ie (modific
re
unor
cte
norm
tive, st
bilire
unor reguli gener
le de ev
lu
re noi etc.).
Principiul
presupune c unit
te
p
trimoni
l i continu n mod norm
l
continuit ii func ion
re
ntr-un viitor previzibil, fr
fi n st
re de lichid
re

ctivit ii s
u de reducere sensibil

ctivit ii;

tunci cnd func ion
re
este delimit
t n timp sunt men ion
te
d
tele de ncepere i de ncet
re

ctivit ii;
permite fix
re
respons
bilit ii entit ii n r
port cu ter e
perso
ne, inclusiv cu bugetul st
tului.
Principiul
presupune delimit
re
n timp
cheltuielilor i veniturilor
independen ei
ferente
ctivit ii unit ii p
trimoni
le pe msur

ng
jrii
exerci iului
cestor
i trecerii
cestor
l
rezult
tul exerci iului l
c
re se
refer;
independen
exerci iului trebuie n ele
s prin
cee
c nu se m
i
po
te interveni
supr
bil
n ului dup perio
d
de r
port
re
(
prob
re
consiliului de
dministr
ie, nregistr
re
l
org
nele
fin
nci
re etc.).
Principiul
int
ngibilit ii
bil
n ul de deschidere
unui exerci iu trebuie s corespund cu
bil
n ul de nchidere
l exerci iului precedent.
bil
n ului de
deschidere
22Principiul
necompensrii
elementele de
ctiv i de p
siv trebuie s fie ev
lu
te i
nregistr
te n cont
bilit
te sep
r
t, nefiind
dmise compens
re

ntre posturile de
ctiv i de p
siv
le bil
ului;
necompens
re
trebuie n ele
s n sensul c pentru fiec
re
element p
trimoni
l cu subst
m
teri
l, pentru orice resurs
c
re reflect drepturi i oblig
ii trebuie s fie deschis, n
cont
bilit
te cte un cont nou;
necompens
re
nu trebuie confund
t cu rectific
re
prevzut n
scopul reflectrii v
lorii rm
se;
v
lorile rectific
tive sunt eviden i
te distinct chi
r d
c n c
zul
unor elemente p
trimoni
le nregistr
re
n bil
n se f
ce l

v
lo
re
consider
t net s
u rm
s (imobilizri rectific
te pe
c
le

mortizrii, m
teri
le de n
tur
obiectelor de invent
r
rectific
te pe c
le
uzurii s
u deprecierilor rectific
te pe se
m

provizio
nelor etc.).
2.5. Func iile cont
bilit ii
Cont
bilit
te

fin
nci
r

Func i
de nregistr
re complet
tr
nz
c iilor entit ii n scopul

determinrii periodice
situ
iei p
trimoni
le i
rezult
tului glob
l;
Func i
de comunic
re fin
nci
r extern (inform
re
ter ilor);
Func i
de instrument de verific
re i de prob judici
r i fisc
l;
Func i
de instrument de gestiune
entit ii;
Func i
de furniz
re
inform
iilor neces
re re
lizrii sintezelor
m
croeconomice;
Func i
de inform
re pentru
n
lize fin
nci
re.
Cont
bilit
te

intern de gestiune
Func i
de determin
re
costurilor pe produse, lucrri i secto
re
de
ctivit
te;
Func i
de determin
re
diferitelor m
rje i
rezult
telor

n
litice pe produse i
ctivit i;
Func i
de gener
liz
re i furniz
re
inform
iilor destin
te
el
borrii bugetelor i costurilor previzion
te;
23 Func i
de gener
liz
re i furniz
re
inform
iilor destin
te

ctu
lizrii indic
torilor de gestiune din structur
t
bloului de
bord
l entit ii;
Func i
de gener
liz
re i furniz
re
inform
iilor destin
te
msurrii perform
n elor l
nivelul secto
relor i pe produse,
lucrri i servicii.
Dup p
rcurgere

cestei uniti de nv
re trebuie s reinei:
C
re este obiectul de studiu
l cont
bilit ii.
C
re sunt procedeele specifice metodei cont
bilit ii.
C
re sunt principiile cont
bilit ii.
C
re sunt func iile cont
bilit ii i necesit
te

plicrii
cestor

n cont
bilit
te
fin
nci
r i n cont
bilit
te
de gestiune.
24UNITATEA DE NV ARE 3
ACTIVUL PATRIMONIAL
3.1. Obiective
L
sfritul
cestei uniti de nv
re studenii vor fi c
p
bili s:
define
sc no iune
de
ctive p
trimoni
le;
identifice i s descrie elementele componente
le
ctivelor
imobiliz
te;
identifice i s descrie elementele componente
le
ctivelor
circul
nte.
3.2. Activul p
trimoni
l
Este form
t din tot
lit
te
mijlo
celor economice destin
te bunei desfurri


ctivitii. Elementele p
trimoni
le de
ctiv se pot cl
sific
: dup modul de v
lorifi
c
re i dup
gr
dul de lichidit
te. Se observ c, dup
mbele criterii se conture
z dou m
ri c
tegor
ii de

ctive p
trimoni
le:
ctive imobiliz
te i
ctive circul
nte.
I. Activele imobiliz
te
-
ctive fixe ce se consum i se v
lorific n mod trept
t, pe
p
rcursul m
i multor cicluri de explo
t
re,
vnd un gr
d de
lichidit
te sczut.
n func ie de n
tur
lor, ele cuprind urmto
rele grupe:
1) Active imobiliz
te
necorpor
le
2) Active imobiliz
te
- sunt
ctive fixe nem
teri
le, reprezent
te prin documente juridice
s
u comerci
le ce confer entit ii
numite drepturi.
- sunt
ctive fixe m
teri
le.
corpor
le
3) Active imobiliz
te
fin
nci
re
- sunt
ctive fixe de n
tur
investiiilor fin
nci
re n c
drul
ltor

ntreprinderi, efectu
te pe termen mediu i lung, n scopul

obinerii de venituri.
1) Activele imobiliz
te necorpor
le cuprind:
) Cheltuielile de
constituire
- sunt cheltuielile efectu
te de entit
te l
nfiin
re
ei: cheltuieli
de nscriere l
Registrul Comerului, t
xe de public
re l

Monitorul Ofici
l, cheltuieli cu emisiune
de
ciuni i pri
soci
le etc.
b) Cheltuielile de
cercet
re-dezvolt
re
- sunt cheltuieli efectu
te pentru
chiziion
re
s
u re
liz
re
n
producie proprie
unor proiecte de cercet
re ce urme
z
fi

plic
te n procesul de producie, n scopul obinerii de profit.
c) Concesiunile
- sunt convenii prin c
re entit
te
dobndete, n schimbul unei
sume, dreptul de
explo
t
pe o
numit perio
d, bunuri
propriet
te
st
tului (terenuri, cldiri etc.).
d) Brevetele
- sunt documentele prin c
re i se recuno
te unei perso
ne dreptul
de
explo
t
exclusiv un produs
l crui
utor este.
e) Licenele de
f
bric
ie
f) Mrcile de f
bric
- sunt drepturi ctig
te de entit
te, de
explo
t
un brevet, prin
cumpr
re

cestui
.
- sunt sume investite de entit
te pentru c
produsele s
le s se
deosebe
sc de produse simil
re existente pe pi
.
g) Fondul comerci
l
- reprezint dreptul supliment
r cuvenit entit ii, peste v
lo
re

bunurilor m
teri
le, d
torit existenei unor condiii deosebite
precum: v
dul comerci
l, clientel
etc.
h) Progr
mele
- reprezint p
rte
soft
echip
mentului electronic, ce permite
inform
tice efectu
re
diverselor oper
iuni cu
jutorul tehnicii de c
lcul.
i) Alte imobilizri - cuprind cheltuieli oc
zion
te de
chiziion
re
s
u el
bor
re

necorpor
le
j) Imobilizrile
necorpor
le n curs
progr
melor inform
tice i
lte imobilizri necorpor
le.
- reprezint costul de producie s
u de
chiziie
imobilizrilor
necorpor
le netermin
te pn l
sfritul exerciiului fin
nci
r.
D
torit utilizrii lor pe dur
t ndelung
t, imobilizrile necorpor
le se supun uzurii n
timp. Expresi
v
loric
uzurii imobilizrilor se nregistre
z n cont
bilit
te sub form



mortizrii.
2) Activele imobiliz
te corpor
le cuprind:

) Terenurile
- n func ie de destin
ie se cl
sific n: terenuri
gri
cell surf
ce receptors. Their effects
re direct

nd very r
pid. With lig
nd-
ctiv
ted (or lig
nd-g
ted) ion ch
nnels (1), binding of the lig
nd to the receptor ch
nges the conform
tion of
the receptor protein. This c
uses
n ion-specific
ch
nnel in the receptor protein to open. The resulting flow of ions ch
nges the electric ch
rge
of the cell membr
ne. Receptors with lig
nd274
Genetics
nd Medicine
Genetic Defects in Ion Ch
nnels
More th
n 20 different disorders due to defec-

tive ion ch
nnel proteins resulting from gene

mut
tions
re known. Such disorders include
cystic fibrosis (see p. 276), the long-QT syndrome,
speci
l type of de
fness, heredit
ry
hypertension (Liddle syndrome), f
mili
l persist
nt hyperinsulinemic hypoglycemi
of inf
ncy, some heredit
ry muscle dise
ses,
nd
m
lign
nt hyperthermi
(see p. 372),
mong
other disorders.
A. Long-QT syndrome,
genetic
c
rdi
c
rrhythmi

Congenit
l long-QT syndrome is ch
r
cterized
by
prolonged QT interv
l in the electroc
rdiogr
m (more th
n 460 ms, corrected for he
rt
r
te), sudden
tt
cks of missed he
rt be
ts
(syncopes) or series of r
pid he
rt be
ts (tors
des de pointes),
nd
n incre
sed risk for sudden de
th from ventricul
r fibrill
tion in
children
nd young
dults.
B. Different molecul
r types of
long-QT syndrome
Prolong
tion of the QT interv
l in the electroc
rdiogr
m results from
n incre
se in the dur
tion of the c
rdi
c
ction potenti
l (1). The
norm
l potenti
l l
sts
bout 300 ms (ph
ses 1

nd 2). The resting membr
ne potenti
l (ph
se
3) is re
ched by progressive in
ctiv
tion of c
lcium currents
nd incre
sing depletion of
pot
ssium currents, which repol
rize the cell.
In ph
se 0 the cell is quickly depol
rized by
ctiv
ted sodium currents following
n excit
tory
stimulus.
LQT1
ccounts for
bout h
lf of the p
tients
with long-QT syndrome. The gene for LQT2 encodes
1195-
mino-
cid tr
nsmembr
ne protein responsible for the other m
jor pot
ssium
ch
nnel th
t p
rticip
tes in ph
se 3 repol
riz
tion (HERG st
nds for (hum
n-ether-r-go-gorel
ted gene,
Drosophil
homologue). LQT3,

sodium ch
nnel protein, consists of four subunits, e
ch cont
ining six tr
nsmembr
ne
dom
ins
nd
number of phosph
te-binding
sites. Homozygosity for LQT1 (KVLQT1 gene) or
LQT5 (KCNE1 gene) c
uses
form of long-QT
syndrome
ssoci
ted with de
fness, the Jervell

nd L
nge-Nielsen syndrome. (Figure
d
pted
from Ackerm
n
nd Cl
ph
m, 1997.) It is import
nt to distinguish the different types bec
use
the choice of medic
tion differs.
References
Ackerm
n, M.J., Cl
ph
m, D.D.: Ion ch
nnels
B
sic science
nd clinic
l dise
se. New Eng.
J. Med. 336 : 1575 1586, 1997.
Ke
ting, M.T., S
nguinetti, M.C.: Molecul
r
genetic insights into c
rdiov
scul
r dise
se.
Science 272 :681 685, 1996.
Schulze-B
hr, E., et
l.: KCNE1 mut
tions c
use
Jervell
nd L
nge-Nielsen syndrome. N
ture
Genet. 17 :267 268, 1997.
Schulze-B
hr, E., et
l.: The long-QT syndrome.

Current st
tus of molecul
r mech
nisms. Z.

K
rdiol. 88 :245 254, 1999.
Viskin, S. : Long QT syndromes
nd tors
des de
pointes. L
ncet 354 :1625 1633, 1999.
Ex
mples of Dise
ses due to Genetic Ion Ch
nnel Defects
Dise
se Inherit
nce Ion Ch
nnel Gene
Locus
Cystic fibrosis AR CFTR (chloride) 7q31
Long-QT syndrome (6 types) AD KVLQT1 (c
rdi
c pot
ssium) 11p15.5
HERG 7q35 36
SCNA5 (c
rdi
c sodium) 3p21
Three other types (see text)
M
lign
nt hyperthermi

AD
AD,
utosom
l domin
nt; AR,
utosom
l recessive.
(Ry
nodine, muscle c
lcium) 19q13.1
(skelet
l-muscle sodium) 17q23 25
(About 18 others not listed,
d
pted from Ackerm
n
nd Cl
ph
m, 1997.)
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.275
Genetic Defects in Ion Ch
nnels
Rom
no W
rd syndrome
(Long-QT syndrome)
Prolonged QT interv
l in
the electroc
rdiogr
m
Syncope
Prolonged QT
Sudden de
th
Autosom
l domin
nt
Six genes involved
(LQT1 - LQT6)
Tors
de de pointes
Type Locus Gene
LQT1 11p15.5 KCNQ1 (KVLQT1)
LQT2 7q35-36 HERG
LQT3 3p21-24 SCNA5
LQT4 4q25-27 unknown
LQT5 21q22.1 KCNE1
LQT6 21q21.1 KCNE2
Long-QT
nd De
fness (Jervell
nd
L
nge-Nielsen) due to
llelic mut
tions

t LQT1
nd LQT5
(
utosom
l recessive)
1. M
in fe
tures
2. Electroc
rdiogr
m
3. Genetics
A. Long-QT syndrome,
genetic c
rdi
c
rrhythmi

+47mV
1
Prolonged c
rdi
c

ction potenti
l
2
3
0
Current
cl
mp
4
85mV
Norm
l

0
100
200
300
400
500
Milliseconds
1. Incre
sed dur
tion of c
rdi
c
ction potenti
l
LQT3 (3q21-24)
SCN5A=N

II
III
LQT1 (11p15.5)
"
IV
Cell membr
ne
C
581
N
1
!KPQ
P
N
1
KvLQT1=IKs
C
2016
2. Volt
ge-
ctiv
ted K-ch
nnel del
yed
in ph
se 3
"
LQT2 (7q35-36)
P
P
HERG=IKr
P
P
4. N
-ch
nnel f
ils to in
ctiv
te completely
during ph
se 0
B. Different molecul
r types of long-QT syndrome
N
1
C
1159
3. Volt
ge-g
ted K-ch
nnel del
yed
in ph
se 3
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.276
Genetics
nd Medicine
Chloride Ch
nnel Defects:
Cystic Fibrosis
Cystic fibrosis (mucoviscidosis) is
highly v
ri
ble multisystemic disorder due to mut
tions of
the cystic fibrosis tr
nsmembr
ne conduction
regul
tor gene (CFTR). Cystic fibrosis (CF) is one
of the most frequent
utosom
l recessive
heredit
ry dise
ses in popul
tions of Europe
n
origin (
bout 1 in 2500 newborns). The high
frequency of heterozygotes (1 : 25) is thought to
result from their selective
dv
nt
ge due to reduced li
bility to epidemic di
rrhe
(choler
).

A. Cystic fibrosis: clinic
l
spects

The dise
se prim
rily
ffects the bronchi
l system
nd the g
strointestin
l tr
ct. Viscous
mucus form
tion le
ding to frequent, recurrent
bronchopulmonic infections
nd eventu
lly
chronic oxygen deficiency ch
r
cterize the
common, severe form of the dise
se. The
ver
ge life expect
ncy in typic
l CF is
bout 30
ye
rs. The dise
se m
y t
ke
less severe,
lmost
mild course. Congenit
l bil
ter
l
bsence of the
v
s deferens (CBAVD) occurs in 95% of p
tients
with CF. It m
y be the only m
nifest
tion in individu
ls with different mut
nt
llelic combin
tions
t the CF locus.
B. Position
l cloning of the gene for
cystic fibrosis (CF)
The CFTR gene w
s isol
ted on the b
sis of its
chromosom
l loc
tion (position
l cloning) on
the long
rm of chromosome 7
t q31 (7q31).
Since the gene could be m
pped to the long
rm
of chromosome 7 ne
r
m
rker locus D7S15,

long-r
nge restriction m
p comprising
bout
1500 kb cont
ining the presumptive CF locus
fl
nked by two m
rker loci, MET
nd D7S8, w
s
constructed. From there
region of 250 kb w
s
isol
ted by
combin
tion of chromosome
w
lking
nd chromosome jumping. Sever
l
genes were loc
ted in this region (c
ndid
te
genes) between the m
rker loci D7S340
nd
D7S424. The gene sought w
s identified by the
finding of mut
tions of this gene in p
tients but
not in controls, by comp
ring with simil
r
genes in other org
nisms (evolution
ry conserv
tion), by determining its exon/intron structure, by sequencing it,
nd by determining the
expression p
ttern of the gene in different tissues.
C. The CFTR gene
nd its protein
The CFTR gene is l
rge, extending over 250 kb of
genomic DNA,
nd is org
nized into 27 exons
(24
re shown in the di
gr
m) encoding
6.5 kb
tr
nscript with sever
l
ltern
tively spliced
forms of mRNA. The CFTR protein h
s 1480

mino
cids. It is
membr
ne-bound chloride
ion ch
nnel regul
tor with sever
l function
l
dom
ins: two nucleotide-binding dom
ins (encoded by exons 9 12
nd 19 23),
regul
tory
dom
in (exons 12 14
),
nd two tr
nsmembr
ne-sp
nning dom
ins (exons 3 7
nd 14b
18). E
ch of the two tr
nsmembr
ne regions
consists of six tr
nsmembr
ne segments. The
nucleotide-binding dom
in 1 (NBD1) confers
cAMP-regul
ted chloride ch
nnel
ctivity. The
most common mut
tion (occurring in 66% of
p
tients),
deletion of
phenyl
l
nine codon in
position 508 ( F508), is loc
ted here. The protein is
member of the ATP-binding c
ssette
(ABC) f
mily of tr
nsporters. The R dom
in cont
ins put
tive sites for protein kin
se A
nd protein kin
se C phosphoryl
tion. CFTR is widely

expressed in epitheli
l cells. The more th
n 800

different mut
tions observed in the CFTR gene
(see http://www.genet.sickkids.on.c
/cftr) c
n be
grouped into five different function
l cl
sses:
(i)
bolished synthesis of full-length protein, (ii)
block in protein processing, (iii) reduced
chloride ch
nnel regul
tion, (iv) reduced
chloride ch
nnel conduct
nce, (v) reduced

mount of norm
l CFTR protein. The underlying
genetic defects include missense mut
tions,
nonsense mut
tions, RNA splicing mut
tions,

nd deletions. Aside from F508 in
bout 66%
of p
tients, the most frequent mut
tions worldwide
re G542X (2.4%), G551D (1.6%), N1303K
(1.3%),
nd W1282X (1.2%).
References
Chillon, M., et
l.: Mut
tions in the cystic fibrosis gene in p
tients with congenit
l
bsence
of the v
s deferens. N. Engl. J. Med.
332 :1475 1480, 1995.
Rosenbluth, D.B., Brody, S. L.: Cystic fibrosis, pp.
329 338, In: J.L. J
meson, ed., Principles of
Molecul
r Medicine. Hum
n Press, Totow
,
N.J., 1998.
Rosenstein, B.J., Zeitline, P.C.: Cystic fibrosis.
L
ncet 351 :277 282, 1998.
Tsui, L.C.: The spectrum of cystic fibrosis mut
tions. Trends Genet. 8 :392 398, 1992.
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.Chloride C
h
nnel Defects: Cystic Fibrosis
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.
277278
Genetics
nd Medicine
Rhodopsin,
Photoreceptor
The hum
n retin
cont
ins
bout 110 million
rod cells for vision in the d
rk
nd 6 million
cone cells for color vision in the light. These
cells cont
in photoreceptors th
t convert light
into
nerve impulse. Rhodopsin is the photoreceptor for we
k light. The light-tr
nsmitting
system consists of numerous components
coded for by genes th
t
re simil
r in structure

nd function to genes for other tr
nsmembr
ne
sign
l-tr
nsmitting molecules.
A. Rod cells
A rod cell consists of
n outer segment with

photoreceptor region
nd
n inner segment
comprising cell nucleus
nd cytopl
sm with endopl
smic reticulum, Golgi
pp
r
tus,
nd mitochondri
. The outer segment cont
ins
bout
1000 disks with rhodopsin molecules in the
membr
ne. In the periphery, the
pproxim
tely
16 nm thick disks
re folded by the protein peripherin. (Di
gr
m
fter Stryer, 1995).
B. Photo
ctiv
tion
In 1958, George W
ld
nd co-workers discovered th
t light isomerizes 11-cis-retin
l (1)
very r
pidly into
ll-tr
ns-retin
l (2),
form

th
t pr
ctic
lly does not exist in the d
rk

(!1 molecule/1000 ye
rs). The light-induced
structur
l ch
nge is so gre
t th
t the resulting

tomic motion c
n trigger
reli
ble
nd reproducible nerve impulse. The
bsorption spectrum of rhodopsin (3) corresponds to the spectrum of sunlight, with
n optimum
t

w
velength of 500 nm. Although vertebr
tes,

rthropods,
nd mollusks h
ve
n
tomic
lly
quite different types of eyes,
ll three phyl
use
11-cis-retin
l for photo
ctiv
tion.
D. Rhodopsin
Rhodopsin is
seven-helix tr
nsmembr
ne
protein with binding sites for function
lly import
nt molecules such
s tr
nsducin, rhodopsin kin
se,
nd
rrestin on the cytosol side. The
binding site of the light-sensitive molecule
(chromophore) is lysine in position 296 of the
seventh tr
nsmembr
ne dom
in. The light-
bsorbing group consists of 11-cis-retin
l. The

mino end of rhodopsin is loc
ted in the disk intersp
ces,
nd the c
rboxy end on the cytosol
side. About h
lf of the molecule is cont
ined in
the seven tr
nsmembr
ne hydrophobic dom
ins, one-fourth in the disk intersp
ces
nd
one-fourth on the cytosol side.
E. cGMP
s tr
nsmitter in the
vizu
liz
tion process
The light c
sc
de ends with r
pid hydrolysis of
cGMP, the intern
l tr
nsmitter in visu
liz
tion.
This le
ds to r
pid closure of the sodium ion
ch
nnels
nd hyperpol
riz
tion of the membr
ne to initi
te nerve impulse, which is tr
nsmitted
s
sign
l to the br
in.
References
P
lczewski, K. et
l.: Cryst
l structure of
rhodopsin: A G protein-coupled receptor.
Science 289 :739 745, 2000.
Schoenlein, R.W., et
l.: The first step in vision:
Femtosecond isomeriz
tion of rhodopsin.
Science 254 :412 415, 1991.
Stryer, L.: Biochemistry. 4 th ed. W.H. Freem
n,
New York, 1995.
Stryer, L.: Molecul
r b
sis of visu
l excit
tion.
Cold Spring H
rbor Symp Qu
nt Biol. 53 :28
294, 1988.
C. Light c
sc
de
Photo
ctiv
ted rhodopsin triggers
series of
enzym
tic steps (light c
sc
de). First,
sign
ltr
nsmitting protein of visu
liz
tion, tr
nsducin, is
ctiv
ted by photo
ctiv
ted rhodopsin.
Tr
nsducin belongs to the G protein f
mily, i.e.,
it c
n
ssume
n in
ctive GDP
nd
n
ctive GTP
form. GTP
ctiv
tes phosphodiester
se. This
very r
pidly hydrolyzes cGMP
nd lowers the
cGMP concentr
tion in cytosol, which le
ds to
closure of the sodium ion ch
nnels. Immedi
tely there
fter, phosphodiester
se is in
ctiv
ted by me
ns of
G protein cycle.
P
ss
rge, Color Atl
s of Genetics 2001 Thieme

All rights reserved. Us
ge subject to terms
nd conditions of license.Rhodopsin,

Photoreceptor
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
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nd conditions of license.
279280
Genetics
nd Medicine
Mut
tions in Rhodopsin
Retinitis pigmentos
(RP) is
genetic
lly heterogeneous group of dise
ses th
t le
d to
pigment
l degener
tion of the retin

nd progressive blindness. Numerous mut
tions in the
rhodopsin gene h
ve been shown to be the
c
use of different forms of RP. Mut
tions in
other genes coding for proteins of the light c
sc
de m
y
lso c
use retinitis pigmentos
.
A. Retinitis pigmentos

The fundus of the eye shows distinct displ
cement of pigment
tion, with irregul
r hyperpigment
tion
nd depigment
tion. The p
pill

(optic disk) shows w
xy yellow discolor
tion.
The loss of vision, especi
lly in dim light (night
blindness), proceeds from the periphery to the
center
t different r
tes depending on the form
of the dise
se, until only
very n
rrow centr
l
visu
l field rem
ins. (Photogr
ph from E. Zrenner, Tbingen.)
B. Point mut
tion in codon 23
The first point mut
tion demonstr
ted in the
rhodopsin gene (Dryj
et
l., 1990) w
s
tr
nsversion from cytosine to
denine in codon
23. This ch
nged the codon CCC for proline (Pro)
into CAC for histidine (His). Since the proline in
position 23 occurs in more th
n ten rel
ted G
protein receptors, it must be very import
nt for
norm
l function.
C. Mut
tions in rhodopsin
The gene locus for rhodopsin (RHO) in m
n lies
on the long
rm of chromosome 3 in region 2,
b
nd 1.4 (3q21.4). Domin
nt
nd
utosom
l recessive inherited mut
tions h
ve been demonstr
ted in hum
ns. Most mut
tions le
d to the
exch
nge of
n
mino
cid,
lthough deletions
m
y
lso occur. Of the 348
mino
cids of
rhodopsin, 38
re identic
l (inv
ri
nt)
t
v
rious positions in vertebr
tes. More th
n 100
different mut
tions
re known for
utosom
l
domin
nt inherited RP. An incre
sing number
of mut
tions
re recognized to c
use
utosom
l
recessive RP. In
ddition, mut
tions in sever
l
other gene loci h
ve been recognized to le
d to
retinitis pigmentos
, e. g. mut
tions in the gene
for peripherin on the short
rm of chromosome
6 in hum
ns (6p)
nd
locus in the centromeric
region of chromosome 8. Other photoreceptor
gene dise
se loci
re the
nd suunits of
phosphodiesterase (PDE).
D. Demonstration of a mutation in
codon 23 y means of
oligonucleotides after PCR
This pedigree (1) with autosomal dominant in-

herited retinitis pigmentosa due to mutation in

codon 23 includes 13 affected individuals in
three generations (affected females, lack
circles; affected males, lack squares). Using
polymerase chain reaction (PCR) (see p. 166),
Dryja et al. (1990) demonstrated the mutation
in amplified fragments of exon 1 (2). The normal oligonucleotide corresponds to the normal
sequence etween codons 26 and 20. The mutant sequence of the oligomere RP contains the
mutant sequence CAC. All affected individuals
gave a hyridization signal with the RP oligomer (2) (II-2, II-12, and III-4 were not examined), whereas unaffected individuals did
not (see p. 408 for demonstration of a point mutation with oligonucleotides).
References
Barkur, S. S. : Retinitis pigmentosa and related
disorders. Am J Med Genet. 52 :467 474,
1994.
Dryja, T.P.: Retinitis pigmentosa, pp. 4297
4309, In: C.R. Scriver, et al., eds., The Metaolic and Molecular Bases of Inherited Disease. 7 th ed. McGraw-Hill, New York, 1995.
Dryja, T.P., et al.: A point mutation of the
rhodopsin gene in one form of retinitis pigmentosa. Nature 343 :364 366, 1990.
McInnes, R.R., Bascom, R.A.: Retinal genetics: a
nullifying effect for rhodopsin. Nature
Genetics 1 :155 157, 1992.
Wright, A.F.: New insights into genetic eye disease. Trends Genet. 8 :85 91, 1992.
Passarge, Color Atlas of Genetics 2001 Thieme
All rights reserved. Usage suject to terms and conditions of license.Mutations
in Rhodopsin
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281282
Genetics and Medicine
Color Vision
As suggested y Thomas Young in 1802, color
vision in humans is mediated y three receptor
types in the cone cells of the retina, one each for
lue, green, and red.
A. Genes for photoreceptor proteins in
cones
The gene for the lue receptor is autosomal; the
genes for the red and green receptors are X
chromosomal. The asorption spectra of the
three receptors show maxima of 426 nm for
lue, aout 530 for green, and aout 550 for red.
The red receptor was discovered to e polymorphic, with two somewhat different asorption
maxima at 552 and 557 nm.
B. Evolution of the genes for visual
pigment photoreceptors
The photoreceptor genes arose from a single ancestral gene (protogene). The rhodopsintransducin pair is found in inverterates and is at
least 700 million years old. The lue receptor is

almost as old as rhodopsin, aout 500 million

years. The separation into a receptor for green
and one for red must have occurred only aout
30 million years ago, after the Old World and
New World apes separated, since man and the
Old World apes have three cone pigments
whereas New World apes have two.
C. Structural similarity of the visual
pigments
In 1986, J. Nathans and co-workers sequenced
the genes for color photoreceptors and oserved marked structural similarities, especially of the green and red receptor genes.
Here the gene products (the receptors) are
shown and their similarities are compared. The
dark dots indicate variant amino acids; the light
dots are identical amino acids, given in percentages.
D. Polymorphism in the photoreceptor
for red
A. G. Motulsky and co-workers (Winderickx et
al., 1992) demonstrated variant codons in three
regions of the red receptor gene (1). Serine was
found at position 180 in 60% of the investigated
males; alanine in 40%. Position 230 showed
polymorphism of isoleucine (Ile) and threonine
(Thr); position 233 of alanine (Ala) and serine
(Ser) (2). Differences in red color perception
could e demonstrated y the color-mixing test
procedure of Raleigh (3).
E. Normal and defective redgreen
vision
One gene for red and one to three genes for
green lie close together on the long arm of the X
chromosome in humans (1). Since the
sequences of these genes are very similar, unequal crossing-over is not infrequent (2). Intergenic crossing-over leads to loss (green lindness) or duplication; intragenic crossing-over
leads to a hyrid gene (red lindness). Green
lindness results from loss of a gene for the
green receptor; red lindness, from a defective
or asent red receptor. With redgreen lindness, neither a normal red nor a normal green
receptor is present. Aout 1% of all men are red
green lind and aout 2% are green lind. Aout
8% show weakness in differentiating red from
green.
References
Kohl, S. , et al.: Total colour lindness is caused
y mutations in the gene encoding the subunit of the cone photoreceptor cGMPg
ted c
tion ch
nnel. N
ture Genet. 19 :257
259, 1998.
Motulsky, A.G., Deeb, S. S.: Color vision
nd its
genetic defects, pp. 4275 4295. In: C.R.
Scriver, et
l., eds., The Met
bolic
nd
Molecul
r B
ses of Inherited Dise
se. 7 th ed.
McGr
w-Hill, New York, 1995.
N
th
ns, J., Thom
s, D., Hogness, D.S. : Molecu-

l
r genetics of hum
n color vision: the

genes encoding blue, green,
nd red pigments. Science 232 :193 202, 1986.
Neitz, M., Neitz, J.: Numbers
nd r
tios of visu
l
pigment genes for norm
l red-green color
vision. Science 267 :1013 1016, 1995.
Winderickx, J., et
l.: Polymorphism in red photopigment underlies v
ri
tion in colour
m
tching. N
ture 356 :431 433, 1992.
Wissinger, B., Sh
rpe, L.T.: New
spects of
n old
theme: The genetic b
sis of hum
n color vision. Am. J. Hum. Genet. 63 :1257 1262,
1998.
Wissinger, B., et
l.: Hum
n rod monochrom
cy: link
ge
n
lysis
nd m
pping of

cone photoreceptor expressed c
ndid
te
gene on chromosome 2q11. Genomics
51 : 325 331, 1998.
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.Color Visi
on
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.
283284
Genetics
nd Medicine
He
ring
nd De
fness
Acoustic sign
ls
re essenti
l for
n
nim
ls
aility to respond appropriately to its environment. Hearing is orchestrated y a large ensemle of proteins acting in concert. Specialized sensory cells in the cochlea of the inner ear
process the incoming sound waves, converting
them into cellular information that is relayed to
the rain via the acoustic nerve. A missing or
defective protein involved in the hearing
process results in hearing loss. Hearing loss is
common in humans. One out of 1000 neworns
lacks the aility to hear or has severely impaired
hearing. Two categories of genetic hearing loss
can e distinguished: nonsyndromic and syndromic. In the former category the genetic defect is limited to the ear; in the latter the ear is
one of several organ systems affected.
The types of genes implicated when defective as
the cause of nonsyndromic hearing loss include
those encoding proteins involved in cytoskeletal structure, transcription factors, ion
channels (potassium channel), and intercellular
gap channels composed of junction connexins.
A. The main components of the ear
The auditory system consists of the outer ear,
the middle ear, and the inner ear. Sound waves
are funneled through the outer ear (auricle) and
transmitted through the external ear canal to
the tympanic memrane, which they cause to
virate. These virations are transmitted
through the tympanic cavity of the middle ear
y a chain of three movale ones, the malleus,
the incus, and the stapes. Three major cavities
form the inner ear: the vestiule, the cochlea,

and the semicircular canals. The chochlea is the

site where auditory signals are processed. The
cochlea contains a memranous layrinth filled
with a fluid, the endolymph. The vestiular apparatus includes three semicircular canals
oriented at 90 degree angles to each other.
They respond to rotatory and linear acceleration. Signals received here are transmitted via
the vestiular nerve, which fuses with the
cochlear nerve to form the acoustic nerve. The
latter transmits the information to the rain.
B. The cochlea
The cochlea contains the cochlear duct, which
forms the organ of Corti. The organ of Corti converts sound waves in the endolymph of the
cochlea into intracellular signals. These are
transmitted to auditory regions of the rain.
The organ of Corti contains two types of sensory
cells: one row of inner hair cells and three rows
of outer hair cells. The inner hair cells are pure
receptor cells. Virations induced y sound lead
to slight deflections of the stereocilia and open
potassium channels at the tips of the stereocilia.
The influx of potassium ions at the tips of the
cilia of the hair cells (see C) causes a change in
memrane potential that results in a nerve impulse, which is transmitted as an auditory signal to the auditory cortex of the rain.
Potassium ions are recycled to the supporting
cells and the spiral ligament into the endolymph of the scala media. The tectorial memrane amplifies the sound waves as a resonator.
(Figure adapted and redrawn from Willems,
2000.)
C. The outer hair cell
The outer hair cells comine sensory function
with the aility to elongate and contract when
acoustically stimulated. The apical pole of a hair
cell carries an array of aout 100 cylindrical
stereocilia in a V-shaped arrangement. Each
stereocilium contains an actin molecule, which
enales it to elongate or to contract. The tips of
the stereocilia are connected y tip links. The
potassium channels are formed y the KCNQ4
protein (yellow) and y connexins (red). Important for the structural integrity and dynamics of
the hair cells is a cytoskeleton involving actin,
myosin 7A, myosin 15, and the protein diaphanous. (Figure adapted and redrawn from
Willems, 2000.)
D. Chromosomal locations of human
deafness genes
Almost every human chromosome harors at
least one gene involved in nonsyndromic
monogenic hearing loss. The diagrammatic
presentation shown here is limited to nonsyndromic hearing loss.
References
Roertson N.D., Morton, R.N.D.: Beginning of a
molecular era in hearing and deafness. Clin.

Genet. 55 :149 159, 1999.

Steel, K.P., Bussoli, T.J.: Deafness genes. Expressions of surprise. Trends Genet. 15 :207
211, 1999.

variaz l
diferite grupe de insecte, putnd
junge l
unele specii pn l
50 % (l

l
rvele unor specii de chironomide).
C i r c u l
i
s n g e l u i l
insecte se f
ce cu
jutorul cmruelor inimii i
di
fr
gmelor dors
l i ventr
l. Sub
ciune
muchilor cmruele inimii se dil
t i se
contr
ct consecutiv, i
r sngele este mpins dintr-o cmru n
lt
pn l

ort. n
momentul dil
trii (di
stol
), to
te v
lvulele unei cmrue se deschid i sngele
ptrunde n interior,
tt din sinusul peric
rdi
l ct i din cmru posterio
r. n
momentul contr
ctrii (sistol
) este deschis num
i v
lvul
din f
, prin c
re sngele
este mpins n cmru

nterio
r, i
r celel
lte v
lvule sunt nchise, oprind refluxul
(fig. 24). Frecven
contr
ciilor cmruelor inimii v
ri
z ntre 30 - 140 pe minut, n
funcie de speci
de insect. Sngele
juns n
ort se rev
rs prin orificiul ei
nterior n
c
vit
te
cef
lic i de
colo n tot corpul. Ridic
re
sngelui din sinusul perineur
l n
sinusul viscer
l, i
r din
cest
n cel peric
rdi
l se re
lize
z prin contr
ct
re

ritmic

di
fr
gmelor ventr
l i dors
l, c
re prezint l
ter
l i
nterior sp
ii deschise, prin c

re
sinusurile comunic ntre ele. Sngele
juns n sinusul peric
rdi
l ptrunde din nou n
v
sul dors
l, circulnd din
poi-n
inte i de jos n sus. Org
nele puls
torii
jut n
circul
i
sngelui, mpingndu-l n
ntene, picio
re i
ripi.
Fig. 24 - Schem
funcionrii inimii l
insecte :

- c
mer
inimii n st
re de sistol; b - c
mer
inimii n st
re
de di
stol; Os - ostiole; V
- v
lvule
(dup P o s p e l o v).
4.6. SISTEMUL RESPIRATOR
L
m
re
m
jorit
te
insectelor, l
fel c
l
myri
pode, respir
i
este t r
h e
n . E
se re
lize
z prin intermediul unui sistem de tuburi r
mific
te numite
t r
h e i, c
re comunic cu exteriorul prin nite orificii denumite s t i g m e (f
ig. 25).
Sistemul tr
he
n este c
r
cteristic insectelor terestre; l
insectele
cv
tice (
l
l
rvele de
ephemeroptere, plecoptere, odon
te etc.) respir
i
se f
ce prin br
nhii tr
heene.
S t i g m e l e s
u s p i r
c o l e l e t r
h e e n e (stigm
, spir
cul
) su
nt
orificii
le tr
heelor, situ
te pe prile l
ter
le
le corpului, cte o pereche pe fi
ec
re
segment, lipsind, n gener
l, pe c
p, protor
ce i segmentul
l 9-le

bdomin
l. Numr
ul
29stigmelor v
ri
z l
diferite grupe de insecte (1 - 2 perechi stigme tor
cice i 2
perechi
pe segmentele
bdomin
le l
thys
noptere, o pereche de stigme tor
cice i 6 perech
i

bdomin
le l

noplure etc.).
Fig. 25 - Sistemul tr
he
n l
insecte (Peripl
net
):

- vzut ventr
l; b - vzut dors
l; Tr - tr
hei; St - stigme
(dup M i
l l i D e n n y).
Stigmele sunt mrginite de un c
dru chitinos, ngro
t, numit p e r i t r e m, i
r
orificiul lor continu cu o inv
gin
re sub form de c
mer, denumit
t r i u m (fig.
26). Pereii interni
i c
merei
tri
le sunt cptuii cu periori s
u excrescene
chitino
se, c
re
u rol de filtru. Atriul uneori este form
t din dou c
mere. Desc
hidere

i nchidere
stigmelor se f
ce prin intermediul unor dispozitive speci
le,
cion
te
de
muchii nchiztori i deschiztori.
Fig. 26 - Seciune longitudin
l printr-o stigm
bdomin
l l
furnic (Formic
) :

- stigm deschis; b - stigm nchis; Ds - deschidere
stigmei; C
- c
mer


nterio
r; Cp - c
mer
posterio
r; Mi - muchi nchiztor; Md - muchi

deschiztor; Tr - tr
hee (dup J
n e t).
30T r
h e i l e sunt de origine ectodermic i reprezint inv
ginri
le
tegumentului. Peretele tr
heii este
lctuit din m e m b r
n
b
z
l , un str

t de
celule hipodermice, numit e p i t e l i u s
u m
t r i c e
i i n t i m
, c
re
corespunde cuticulei tegumentului. Cel m
i dezvolt
t nveli cuticul
r este exocutic
ul
,
c
re este ngro
t sub form de spir
l i po
rt denumire
de t e n id i e (tenidium). Epicuticul
i endocuticul
, m
i dezvolt
te n tuburile tr
heene
gro
se, sunt subi
te mult n tr
heele subiri. Tenidi

re rolul de
menine tubul
tr
he
n deschis, de
mpiedic
turtire
lui.
De l
c
mer

tri
l pornete o tr
hee scurt, c
re se r
mific n 3 r
muri : o
r
m u r d o r s
l , c
re deservete inim
, muscul
tur
i tegumentul; o r
m ur
v e n t r
l , c
re merge l
sitemul nervos, muscul
tur i tegument i o r
m u- r v
i s c e r
l spre intestin i celel
lte org
ne viscer
le. R
murile tr
heene, n
continu
re, se divid dicotomic n tr
hei din ce n ce m
i fine i nguste, pn l

dimensiunile c
pil
relor, cu un di
metru m
i mic de un micron, numite t r
h e
o l e
(fig. 27). Aceste
din urm sunt lipsite de tenidie i ptrund n interiorul celulelor
corpului,
vnd pereii perme
bili pentru g
ze i lichide.
L
numero
se insecte (unele hymenoptere, diptere, orthoptere etc.) tuburile
tr
heene longitudin
le prezint dil
tri, m
i
les n regiune

bdomenului, numite
s
c i
e r i e n i, c
re servesc c
rezervo
re de
er n timpul zborului.
Fig. 27 - Schem
tr
heolelor:
Fm - fibr muscul
r; Te - tr
heole;
Tr - tr
hee (dup K e l l e r).
F i z i o l o g i
r e s p i r
i e i. Procesul respir
iei l
insecte se re
liz
e
z
prin
erisire
tr
heelor, respectiv ptrundere

erului n esuturi i elimin
re
bioxid
ului
de c
rbon. Expir
i
este
ctiv, efectundu-se prin
ps
re
sistemului tr
he
n d
torit
contr
ctrii muchilor dorso-ventr
li, c
re
propie tergitele de sternite, i prin
contr
ct
re
muchilor longitudin
li, c
re scurte
z
bdomenul. Inspir
i
, d
torit
revenirii
bdomenului l
norm
l, este p
siv. L
inspir
ie,
erul ptrunde prin stigm
e n
tr
hei i tr
heole. L
nivelul
cestor
din urm se re
lize
z difuziune
oxigenului n
celule.
L
unele insecte (
pterygote) to
te stigmele p
rticip
tt l
inspir
ie ct i l

expir
ie. L

ltele (Acridid
e), unele stigme servesc l
inspir
ie, i
r
ltele l

expir
ie,
nchizndu-se ritmic. Frecven
micrilor respir
torii l
insecte v
ri
z ntre 12 - 100
pe minut, n funcie de specie, f
ctorii mediului
mbi
nt (temper
tur, umidit
te etc.
) i
intensit
te

ctivitii (rep
us, mic
re).
n timpul respir
iei se produce oxid
re
subst
nelor proteice,
grsimilor i

hidr
ilor de c
rbon din esuturi i eliber
re
bioxidului de c
rbon. Re
ci
oxidrii est
e
exotermic i exodin
mic, eliberndu-se energi
i cldur
neces
re ntreinerii
31proceselor vit
le. Elimin
re
bioxidului de c
rbon se f
ce prin tr
hei i num
i n
mic
p
rte (pn l
25 %) este difuz
t prin tegument n mediul exterior.
L
unele insecte inferio
re (
pterygote), c
i l
unele l
rve endop
r
zite (unele
hymenoptere i diptere), respir
i
este cut
nee, oxigenul difuznd prin tegument.
4.7.
SISTEMUL EXCRETOR
n urm
proceselor met
bolice c
re
u loc n esuturi i org
ne, n
f
r

sistemului digestiv, rezult unele subst
ne inerte s
u vtmto
re org
nismului (
cizi
urici, ox
l
i, c
rbon
i etc.), c
re sunt elimin
te sub form de e x c r e t e prin s

istemul
excretor. Funci
de excreie, l
insecte, este ndeplinit de tuburile lui M
lpighi, co
rpul

dipos i
lte org
ne.
T u b u r i l e l u i M
l p i g h i, princip
lul org
n de excreie, sunt de orig
ine
ectodermic. Ele se prezint sub form
unor tuburi lungi i subiri, nchise l
c
petele
libere i deschise l
b
z
lor de fix
re, n
poi
v
lvulei pilorice,
pro
pe de limi
t

intestinului mediu cu cel posterior (v. fig. 22). Numrul tuburilor lui M
lpighi v

ri
z
mult dup grupele de insecte. Astfel, coccidele
u 2 tuburi, thys
nopterele, homop
terele,
dipterele i lepidopterele 4, coleopterele 4 - 6, orthopterele 90 - 120 etc.
Din punct
de vedere histologic tuburile lui M
lpighi sunt form
te dintr-un singur str
t de
celule,
m
ri, epiteli
le, nvelite de membr
n
b
z
l. Tuburile
u o muscul
tur proprie, c
re
le

sigur micrile n c
vit
te
corpului. Din punct de vedere fiziologic, tuburile lui
M
lpighi ndeplinesc rolul de rinichi, extrgnd din snge
cidul uric i diferitele sruri.
Aceste produse de excreie
jung prin c
n
lele tuburilor n intestinul posterior, de
unde
sunt elimin
te prin orificiul
n
l od
t cu excrementele. L
unele insecte, tuburi
le lui
M
lpighi ndeplinesc i
lte funcii c
: org
ne secreto
re de mt
se, c
re servesc l

confecion
rte
coconilor (unele specii de Pl
nipenni
, unele coleoptere etc.) s
u
org
ne secreto
re
unor fermeni. L
fungivoride (Dipter
) p
rte
dors
l
tuburil
or
lui M
lpighi este modific
t n org
ne lumino
se.
C o r p u l
d i p o s s
u c o r p u l g r
s este rspndit n c
vit
te
celomic
sub form

dou str
turi : unul sub tegument - s t r
t u l p
r i e t
l i unu
l n jurul
tubului digestiv - s t r
t u l p e r i v i s c e r
l. Corpul gr
s este
lctui
t dintr-o

glomer
re de celule sferice s
u poliedrice, cu nucleul mic. Culo
re
corpului g
r
s este
v
ri
bil :
lb, verzuie, g
lben, portoc
lie.
Corpul gr
s ndeplinete m
i multe funcii, dintre c
re cele m
i import
nte sunt
urmto
rele : c
org
n excretor i c

cumul
tor de subst
ne de rezerv. Astfel, corpul
gr
s nm
g
zine
z, c
un rinichi de
cumul
re, diferite sruri vtmto
re
org
nismului sub form de crist
le, c
re pot rmne n interiorul lui to
t vi

insectei
s
u sunt elimin
te prin intermediul tuburilor lui M
lpighi. C
subst
ne de rezerv
se

cumule
z n corpul gr
s grsimile, proteinele i m
i
les glicogenul.
C e l u l e l e p e r i c
r d i
l e s
u n e f r o c i t e l e se prezint sub
form

unor m
se celul
re, situ
te n
propiere
v
sului s
nguin, pe fibrele s
u ntre fibr
ele
di
fr
gmei dors
le. Ele extr
g din snge diferite corpuri strine (unele subst
ne
proteice, clorofil
etc.).
G l
n d e l e l
b i
l e se ntlnesc l
unele grupe de insecte inferio
re
(collembole - poduride etc.), l
c
re tuburile lui M
lpighi lipsesc. Excretele l
or sunt
elimin
te printr-un c
n
l c
re se deschide l
b
z
buzei superio
re.
324.8. SISTEMUL SECRETOR
Sistemul secretor
l insectelor este
lctuit din dou tipuri de gl
nde: exocrine i
endocrine. Secreiile gl
ndelor exocrine
jung l
diferite org
ne prin intermediul
unor
c
n
le speci
le. Gl
ndele endocrine sunt lipsite de c
n
le, i
r secreiile lor sun

t
tr
nsport
te cu
jutorul sngelui n to
te regiunile corpului.
Gl
ndele exocrine sunt fo
rte diferite c
origine i funcie. Ele sunt rep
rtiz
te
n diferite regiuni
le corpului insectelor (n tegument, tubul digestiv, org
nele g
enit
le
etc.). Unele dintre gl
ndele exocrine sunt prezent
te n continu
re.
G l
n d e l e c e r i e r e, c
re secret ce
r
, se ntlnesc l
unele grupe de
homoptere (coccide,
leurodide,
fide).
G l
n d e l e l
c c i p
r e se gsesc l
unele specii de coccide (L
ccifer
l
cc
). Ele secret l
curi speci
le, c
re reprezint un
mestec de rini i ce
r.
G l
n d e l e r e p u l s i v e secret subst
ne cu mirosuri puternice i
respingto
re, c
re
u rol de
pr
re. Ele sunt situ
te pe tor
ce i
bdomen (l

Euryg
ster), l

rticul
iile picio
relor (l
Meloe, Coccinell
) etc.
G l
n d e l e p r o d u c t o
r e d e f e r o m o n i, frecvente l
numero
s
e
specii de insecte, sunt form
iuni gl
ndul
re, de o m
re v
riet
te, situ
te n difer
ite
regiuni
le corpului (pe
ripi, pe
bdomen, n
bdomen etc.). Secreiile
cestor gl

nde,
subst
ne chimice prin intermediul cror
indivizii
celee
i specii comunic ntre ei,
sunt cunoscute sub denumire
de f e r o m o n i. Ei sunt n mod tipic odoriferi i

cione
z direct
supr
sistemului nervos
l individului receptor.
n funcie de semnific
i
mes
jului, feromonii sunt mprii n dou m
ri grupe:
feromoni de dezvolt
re (met
bolici) i feromoni de
ciune (de decl
n
re).
F e r o m o n i i d e d e z v o l t
r e, c
re induc l
indivizii receptori une
le
modificri met
bolice s
u de dezvolt
re.
F e r o m o n i d e
c i u n e, c
re
ntrene
z schimbri de comport
ment.
Cele m
i import
nte tipuri sunt red
te m
i jos.
F e r o m o n i d e b
l i z
j, cu c
re este m
rc
t intiner
riul de depl
s
re
spre
surs
de hr
n etc. (l
insectele soci
le, l
c
rii de sco
r i de lemn).
F e r o m o n i d e o v i p o z i i e, cu c
re sunt m
rc
te locurile propice
(n
ri) s
u "interzise" (mutele fructelor) pentru depunere
oulor.
F e r o m o n i d e
l
r m , c
re determin fug
(dispers
re
) indivizilor
popul
iei n momentul
t
cului unui prdtor (l

fide).
F e r o m o n i d e
g r e g
r e, c
re
sigur concentr
re
popul
iei n
vedere
migr
iei s
u populrii unui biotip (lcuste, c
rii de lemn i sco
r) i
coeziune
f
miliei (l
insectele soci
le).
F e r o m o n i s e x u
l i c
re medi
z rel
iile dintre cele dou sexe n
inte,
n timpul i dup mperechere. Sunt cunoscute trei grupe de feromoni sexu
li.
Atr
ct
ni sexu
li, produi de unul din sexe pentru
tr
gere
sexului opus n
vedere
mperecherii. Se cunosc, pn n prezent,
tr
ct
ni sexu
li l
numero
se specii
de insecte din ordinele : Lepidopter
, Coleopter
, Hymenopter
etc. n m
jorit
te

c
zurilor, feromonii sunt produi de femel i
u
ciune
tr
ctiv
supr
m
sculului,
d
r sunt c
zuri cnd feromonul este produs de m
scul i exercit
tr
cie
supr
femelei
.
L
unele specii, reduse c
numr, feromonii sunt produi i de m
scul i de femel.
Atr
ct
nii sexu
li
u o m
re nsemnt
te pr
ctic, utilizndu-se ndeosebi n
prognoz i comb
tere
unor specii de insecte dunto
re.
Afrodisi
ci, produi de ctre m
sculii
tr
i, cu scopul de
excit
femelele i de

le determin
s
ccepte mperechere
.
33Repeleni sexu
li, produi n
p
r
tul genit
l
l m
sculului i introdui od
t cu
sperm
n cel
l femelei. Se m
rche
z
stfel femelele fecund
te i i determin pe
ceil
li m
sculi s le evite.
L
insecte se ntlnesc i
lte tipuri de gl
nde, dintre c
re menionm: g l
n- d e
u r t i c
n t e (l
l
rvele unor specii de lepidoptere: Pythioc
mp
, Lym
ntri

etc.), cu
rol de
pr
re: g l
n d e s e r i c i p
r e, din
cror secreie se forme
z firul
de

mt
se (l
unele l
rve de lepidoptere : Bombyx, Anther
e
etc.); g l
n d e

p r o d u c t o
r e d e v e n i n (l
unele specii de hymenoptere: Apis, Vesp

,
Formic
), c
re servesc l

pr
re etc.
Gl
ndele endocrine. Secreiile
cestor gl
nde, cunoscute sub numele de
h o r m o n i s
u i n c r e t e,
u un m
re rol n vi

insectelor n ce privete
dezvolt
re
l
rv
r, nprlire
, di
p
uz
,
met
morfoz
etc. Au fost studi
te
urmto
rele tipuri de gl
nde endocrine : celulele neurosecretorii
le g
nglionilor
cerebroizi, gl
ndele protor
c
le, corpor

ll
t
i corpor
c
rdi
c
.
C e l u l e l e n e u r o s e c r e t o r i i
l e g
n g l i o n i l o r c e
r e b r oi z i secret h o r m o n u l c r e i e r u l u i. Rolul
cestui hormon se conside
r c este
fo
rte diferit. n unele c
zuri, controle
z i intensific
ctivit
te
gl
ndelor
protor
c
le; n
lte c
zuri determin ncet
re

ctivitii
cestor
din urm l
l
rve i
pupe, frne
z cretere
i dezvolt
re
i determin intr
re
i ieire
din di
p
uz.
Hormonul creierului se po
te
cumul
n corpor
c
rdi
c
i corpor

ll
t
, determinn
d

ctiviz
re

cestor
.
G l
n d e l e p r o t o r
c
l e sunt reprezent
te printr-o pereche de gl
n
de,
situ
te n p
rte
inferio
r
protor
celui, pe l
turile g
nglionilor respectivi, cu
c
re sunt
n legtur. Aceste gl
nde secret h o r m o n u l n p r l i r i i,
m e t
m o r f o z
e i s
u e c d y s o n, c
re determin ntrerupere
di
p
uzei i reglemente
z nprlirile
i, n gener
l, to
t dezvolt
re
l
rv
r.
C o r p o r

l l
t
se prezint sub form

doi lobi, situ
i de
supr
tubului
digestiv
nterior, n p
rte
posterio
r
g
nglionilor cerebroizi. Aceste gl
nde, c

re se
ntlnesc l
l
rvele i l

dulii insectelor superio
re (Pterygot
), precum i l
unele
insecte inferio
re (Apterygot
- Diplur
), secret h o r m o n u l j u v e n i l s

u n e o t
e n i n. Acest hormon intervine n morfogenez i condiione
z cretere
proporion
l

l
rvelor. Reducere
concentr
iei hormonului juvenil n snge determin l
l
rve
tr
nsform
re
n pupe i
poi n insecte
dulte.
C o r p o r
c
r d i
c

p
re sub form

doi lobi, situ
i n p
rte

nterio

r

g
nglionilor cerebroizi. Rolul
cestor gl
nde este m
i puin cunoscut; se consid
er c
ele reglemente
z
ctivit
te
gl
ndelor protor
c
le.
Activit
te
gl
ndelor endocrine se desfo
r n complex, sub controlul direct
l
sistemului nervos centr
l i sub influen
f
ctorilor de mediu.
Cuno
tere
sistemului endocrin i
fiziologiei s
le prezint o m
re v
lo
re
pr
ctic. Pe
ce
st
se b
ze
z comb
tere
biologic
insectelor cu
jutorul produsel
or
hormon
le.
4.9. SISTEMUL NERVOS
Sistemul nervos
l insectelor, c
i
l celorl
lte
rthropode, este de tip
sc
l
riform. El este de origine ectodermic i se compune din sistemul nervos centr

l,
sistemul nervos simp
tic i sistemul nervos periferic.
S i s t e m u l n e r v o s c e n t r
l este
lctuit dintr-un l
n dublu de
g
nglioni, unii ntre ei longitudin
l prin fibre nervo
se numite c o n e c t i v e i
tr
nsvers
l prin c o m i s u r i.
34Sistemul nervos centr
l este constituit din 3 grupe de g
nglioni : g
nglionii
cerebroizi, g
nglionii subesof
gieni i g
nglionii l
nului nervos ventr
l (fig. 28)
.
G
n g l i o n i i c e r e b r o i z i s
u s u p r
e s o f
g i e n i, c
re

forme
z
creierul l
insecte, sunt
ez
i n c
psul
cef
lic, de
supr
esof
gului. Ei sunt
constituii din 3 perechi de g
nglioni: protocerebrum, deutocerebrum i tritocerebru
m.
Protocerebrum, corespunztor primului segment cef
lic (
cronul), reprezint
p
rte
ce
m
i dezvolt
t
creierului i trimite nervi l
org
nele vizu
le (ochii c
ompui
i ocelii).
Deutocerebrum, corespunztor segmentului
nten
l, inerve
z
ntenele, i
r
tritocerebrum, corespunztor segmentului l
br
l, trimite nervi l
buz
superio
r i
regiune
front
l.
Prin inelul periesof
gi
n creierul este leg
t de g
nglionii subesof
gieni.
G
n g l i o n i i s u b e s o f
g i e n i sunt situ
i sub esof
g i s-
u form
t
din
contopire

trei perechi de g
nglioni, c
re corespund segmentelor : m
ndibul
r,
m
xil
r i l
bi
l. Ei inerve
z m
ndibulele, m
xilele i buz
inferio
r.
Fig. 28 - Sistemul nervos l
insecte :

- vzut ventr
l; b - vzut dors
l (g
nglionii cef
lici i tor
cici);
Ant -
ntene; Ao -
ort
; C
ll - corpor

ll
t
; Cc
r - corpor
c
rdi
c
;
Ccen - corp centr
l; Com - comisur; Con - conectiv; Confr - conectiv
front
l; Cped - corp peduncul
t; Deut - deutocerebrum; Ghip - g
nglioni
hipocerebr
li; Gn - g
nglioni; Gs - g
nglioni subesof
gieni; Gven - g
nglioni
ventr
li; N
nt - nervi
nten
li; N
r - nervii
ripilor; Npc - nervii picio
relor
;
Nr - nervul recurent; Nsim - nervii sistemului simp
tic; Oc - ochi; Ocl - oceli;
Oes - esof
g; Pcer - punte
cerebr
l; Prot - protocerebrum; Trit - tritocerebrum
(dup W e b e r).
35L
n u l n e r v o s v e n t r
l este
lctuit, n gener
l, din 3 perechi de
g
nglioni tor
cici i 8 perechi de g
nglioni
bdomin
li. Acest numr de g
nglioni
i
l
nului nervos ventr
l se ntlnete de obicei n st
diul embrion
r i l
unele insecte
inferio
re. L
m
re
m
jorit
te
insectelor numrul g
nglionilor este m
i mic d
t
orit
contopirii
cestor
ntre ei. Astfel, g
nglionii tor
cici fuzione
z
dese
, uneori i
cu
prim
pereche de g
nglioni
bdomin
li, formnd o singur m
s g
nglion
r. De

semene
, ultimele 2 - 3 perechi de g
nglioni
bdomin
li se pot contopi ntre ei. n
r
port cu gr
dul de fuzion
re
g
nglionilor l
insecte se const
t deosebiri evid
ente n
privin
numrului g
nglionilor nervoi,
tt de l
specie l
specie ct i l

cee
i
specie, n funcie de sex s
u st
diul de dezvolt
re. Astfel, l
l
rvele de
lbine ex
ist 3
perechi de g
nglioni tor
cici i 8 perechi de g
nglioni
bdomin
li, pe cnd l

duli
numrul lor se reduce l
2 perechi de g
nglioni tor
cici i 5 perechi
bdomin
li. L

unele insecte se pot contopi toi g
nglionii l
nului nervos ventr
l cu g
nglionii
subesof
gieni ntr-o singur m
s (l
unele homoptere, coleoptere, l
l
rvele unor
diptere etc.). De l
g
nglionii l
nului nervos ventr
l ple
c nervi l
org
nele din
regiunile n c
re se gsesc. Astfel, g
nglionii tor
cici inerve
z muscul
tur

segmentelor respective, picio
rele i
ripile, i
r g
nglionii
bdomin
li muscul
tu
r


bdomenului etc.
S i s t e m u l n e r v o s s i m p
t i c s
u v i s c e r
l este
lctuit din
urmto
rele pri : sistemul nervos stom
tog
stric, nervul ventr
l nepereche i sistemul
simp
tic c
ud
l.
S i s t e m u l s t o m
t o g
s t r i c este n legtur direct cu tritocerebrum i
inerve
z regiune

nterio
r
tubului digestiv (intestinul
nterior i cel mijlociu)
.
N e r v u l v e n t r
l n e p e r e c h e, m
i dezvolt
t n
bdomen, se ntinde su
b
form
unor f
scicule subiri de-
lungul l
nului g
nglion
r ventr
l, printre conect
ivele


cestui
. n fiec
re segment el prezint excrescene l
ter
le, c
re inerve
z stigmele i

tr
heele.
S i s t e m u l s i m p
t i c c
u d
l pornete de l
ultim
pereche de
g
nglioni
bdomin
li i inerve
z regiune
posterio
r
tubului digestiv i org
nele de
reproducere.
S i s t e m u l n e r v o s p e r i f e r i c este situ
t sub hipoderm
tegument
ului.
Este
lctuit din numero
se celule nervo
se, prevzute cu nervi c
re f
c legtur
cu
diferite org
ne i esuturi.
F i z i o l o g i
s i s t e m u l u i n e r v o s. Fiec
re g
nglion este un ce
ntru
reflex. Prin nervii senzitivi, c
re le
g org
nele de sim de lobul respectiv
l
g
nglionului, se primesc impulsuri de l
periferi
corpului, i
r de l
lobul mot
or prin
nervii motori (locomotori) sunt tr
nsmise rspunsurile (comenzile) muchilor org
nul
ui
efector. Prin intermediul
rcului reflex se re
lize
z legtur
ntre diferitele pri
le
corpului i efectu
re
micrilor
cestui
, sistemul nervos centr
l reprezentnd sistemu
l
de legtur i coordon
re
micrilor insectei.
Creierul, c
princip
lul centru
l
ctivitii insectelor, coordone
z funciile
receptorilor optici i olf
ctivi. G
nglionii subesof
gieni dirije
z micrile pieselor
buc
le. G
nglionii cerebroizi i cei subesof
gieni regle
z tonusul muscul
r. Sistem
ul
nervos stom
tog
stric, prin nervul recurent regle
z digesti
i
Willems, P.J.: Genetic c
uses of he
ring loss.
New Engl. J. Med. 342 :1101 1109, 2000.

ctiv
ted protein kin
se (2) further
ctiv
te

substr
te protein. Most protein kin
ses
phosphoryl
te tyrosine (tyrosine kin
se),
serine, or threonine by tr
nsferring
phosph
te
NITATE
C
CONTABI LITATEA FORM
DE BAZ
A EVIDE
EN EI EC ONOMICE
E
1. Obiective e
1.1
L

sfritul
c cestei unit i de nv
r re studenii vor fi c
p
b bili s:
define e
sc no iun
ne
de cont

bilit
te;
n ele

g necesi it
te
org
n nizrii con nt
bilit ii l
nivelul fiecrei
unit i p
trimoni

le;
n ele

g rolul pe c
re l
re cont

bilit
te
n n c
drul
ctivit ii


desf ur
te de un
nit ile p
tri imoni
le;
n ele

g modul de
d org
niz
r re
cont
bil lit ii l
nive el microeco onomic.
1.2. Definir re
, necesit t
te
i rolu
ul cont
bili it ii
C
Cont
bilit
t

te

exist

t din cele e m
i vech hi timpuri. C.G. Dum

mitrescu, n Istori

cont
bil lit ii,
rt

c grecii
u

mprumu ut
t tehnic
eviden ei co ont
bile de l
egipteni, i
r de l

ei
u pr relu
t-o rom
m
nii. D
r se p
re c
eviden ele cont
bile sunt
s
mult m
i
m vechi n n istori

omeniri ii. Cont
bil lit
te
n p

rtid dubl s-
nscu ut c
urm

re
pr
cti cii c
ont
bil lilor din
Vene i

i Genov

. n
nul 1 494, Luc
P
ciolo des scrie cont
b
bilit
te
n p
rtid
p
dub bl ntr-o
lucr
re de m
tem

tic i geom
metrie. Dup
p
p
ri i

cestei luc crri,
plic
r re
cont
bi ilit ii n
p
rtid dubl s-
r spndit i n

lte ri
le Europei.
A
Astzi,
con
nt
bilitii i i revin s
rci ini din ce n n ce m
i gr rele. E
c
u
ut s-i dep pe
sc
limitele , fiind pus n situ
i
de
descrie e org
niz
i ii din ce n ce
c m
i comp
plexe c
re opere
z
o
ntr-un m
mediu econ nomic i soc ci
l n contin
nu mic
re e i tr
nsfor rm
re.
Princip
p
lele
spec cte
c
ins strument de descriere, de
d model
re e
ntreprin nderilor;
sub c
r re trebuie
c
ins strument de prelucr
re
inform
ii ilor;
studi
t cont
bilit t
te
c
pr r
ctic s
u u joc soci i
l, nscri s ntr-o r ree
de
(dup N.
N Fele
g) )
restric cii regleme ent
re m
i mult
m s
u m
i puin stric cte.
10Cont
bilit
te
po
te fi consider
t drept o
rt, o tehnic s
u o tiin, d
r indiferent
de
cum
m privi-o, cont
bilit
te
este un joc soci
l ce
re drept fin
lit
te reprezen
t
re
unei
re
liti c
re este entit
te
.
Cont
bilit
te
studi
z
cele l
turi
le reproduciei soci
le c
re se pot exprim
n
et
lon bnesc. E
urmrete existen
i din
mic
p
trimoniului
genilor economici,
procesele economice, pe c
re
ceti
le org
nize
z, st
bilind i nregistrnd rezult
tele
fin
nci
re fin
le.
1.3. Org
niz
re
evidenei cont
bile l
nivel microeconomic
Conform Legii Cont
bilitii nr. 82/1991, ntreprinderile
u oblig
i
s org
nizeze i
s conduc cont
bilit
te proprie, n limb
romn i n moned
n
ion
l. Org
niz
re

cont
bilitii reprezint deci, nu num
i o necesit
te,

cum
m
rt
t
nterior, d
r i o
oblig
ie impus prin reglementrile leg
le n vigo
re.
De
ltfel, entit
te
c
sistem complex economico-soci
l i
dministr
tiv-org
niz
t
oric,
ndeplinete o serie de funcii, n c
drul cror
un rol eseni
l l
re funci
fin
nci
rcont
bil.
Pentru ndeplinire

cestei funcii i n
cel
i timp pentru respect
re
legii, n c
drul

entit ii se org
nize
z i funcione
z un comp
rtiment speci
liz
t, fin
nci
r-cont
bil. n

c
drul
cestui comp
rtiment lucre
z perso
ne cu studii de speci
lit
te (medii i su
perio
re),

vnd
tribuii distincte n domeniul evidenei cont
bile oper
tive i gener
le.
Comp
rtimentul fin
nci
r-cont
bil se subordone
z cont
bilului ef, c
re
re studii
superio
re
n fin
ne-cont
bilit
te.
Aici se consemne
z zilnic i lun
r, to
te oper
iunile economico-fin
nci
re ce
u loc
n entit
te, respectiv: cumprri, vnzri, consumuri, s
l
rii, nc
sri, pli etc., conduc
fin
l l
determin
re
rezult
tului
ctivitii i l
ntocmire
situ
iilor fin
nci
re
nu

le de
sintez i r
port
re cont
bil.
Lun
c
lend
ristic po
rt denumire
de perio
d de gestiune, i
r
nul c
lend
ristic,
de exerciiu fin
nci
r.
n b
z
d
telor furniz
te de eviden
cont
bil, se pot efectu

n
lize economicofin
nci
re privind corel
re
resurselor
loc
te cu rezult
tele obinute, se pot c

lcul
diveri
indic
tori i se po
te determin
evolui
diverselor fenomene n timp, cu f
ctorii poz
itivi i
neg
tivi c
re le-
u gener
t.
11n c
drul entit ii, cont
bilit
te
se org
nize
z pe dou circuite p
r
lele: cont
bilit

te
fin
nci
r i cont
bilit
te de gestiune.
este reglement
t prin norme unit
re;
ofer o viziune glob
l
supr

ctivitii;

re un obiectiv fin
nci
r reflect
re
im
ginii fidele

p
trimoniului;
Cont
bilit
te
fin
nci
r
genere
z fluxuri de inform
ii i documente externe;

plic reguli norm
tive;
ofer d
te utiliz
torilor externi (furnizori, clieni, bnci,
investitori, org
ne de control etc.);
se refer l
perio
de nchei
te (lun,
n).
se l
s l
l
titudine
fiecrei entit i;
ofer o viziune det
li
t
supr

ctivitii;

re un obiectiv economic constnd n supr
veghere
i
controlul
ctivitii prin intermediul costurilor;
Cont
bilit
te
genere
z fluxuri de inform
ii interne;
de gestiune
plic reguli st
bilite n c
drul entit ii;
ofer d
te conducerii entit ii;
se refer l
prezent i viitor (previziuni);
cl
sific cheltuielile dup locul de re
liz
re i destin
i

lor.
12Dup p
rcurgere

cestei uniti de nv
re trebuie s reinei:
Ce este cont
bilit
te
i c
re este rolul
cestei
n c
drul unei
entiti.
C
re sunt circuitele de org
niz
re
le cont
bilitii n c
drul
unei entiti.
Ce este cont
bilit
te
fin
nci
r i prin ce se c
r
cterize
z.
Ce este cont
bilit
te
fin
nci
r i prin ce se c
r
cterize
z.
13EA DE NV
V ARE 2
UNITATE
OBIEC
CTUL I METODA
M
C
CONTABIL
LIT II

1. Obiective e
2.1
L

sfritul
c cestei unit i de nv
r re studenii vor fi c
p
b bili s:
identi fice obiectu ul de studiu
l cont
bili it ii;
identi fice proced deele ce
p
r r in metode i cont
bilit ii;
identi fice i s n n ele
g rolu ul principiil or cont
bili it ii;
identi fice func ii ile cont
bil lit ii i ne ecesit
te

p
plicrii
ce estor
n
cont
b bilit
te
fin
nci
r i n n cont
bilit

te
de gest tiune.
2 Obiect tul cont
bil lit ii
2.2.
C orice di sciplin tii in ific, con nt
bilit
te
reprezint simult
n
C

s
o teorie
t
i me etod. n
c
lit
te

s
de teor rie tiin ific c, cont
bil it
te
repre ezint un sis stem de pr
i incipii i cu unotin e
c
re ex xplic i inf forme
z, i
r
i c
meto
od s
u teh hnic, un
n ns
mblu co oerent de procedee,
p
instrume ente prin c

re se observ v i nregis stre
z resu ursele econo omice
le so
ociet ii, sep p
r
te c

utilit i p
trimoni
l le.
p ii cu
Concep
1. Concep p i

dmin nistr
tiv consider c obiectu ul cont
bil lit ii l
privire l
constit tuie reflect
re
i con ntrolul, n expresie v
loric,
v

f
ptelor
delimit

re

dmini istr
tive, n n vedere
sp prijinirii m

n
gementu ului, pen
tru
ob ine
i defin
nire
cu un minim de eforturi (ch heltuieli) un n m
xim de e efecte eco onomice
ului
obiectu (rezult t
te-venituri i).
cont
bi ilit ii
2. Concep p i
juridic c consider r c obiec ctul cont
b bilit ii l forme
z
f
p
trim
moniul unui i subiect de
d drept (

gent econo
omic), prin n prism

rel
iil or juridice e,
dic
drepturil lor i obl lig
iilor p pecuni
re
(m
ter ri
le)
le unei
u
perso

ne fizice s
u juridic ce, n core el
ie cu
obiecte ele,
dic cu
u bunurile i
v
lorile co orespunzto o
re.
3. Concep p i

econ omic
s
u u
fin
nci


r
consid
der
c

obiectul
cont
b bilit ii l co
onstituie cir rcuitul c
pit t
lului, priv vit
tt sub
spectul
destin

iei lui, c t i sub for rm
modulu ui de dobn ndire, respe ectiv sub
form
de c
pit
l propriu
p
i c

pit
l strin. .
16 Obiectul cont
bilit ii este reprezent
t de reflect
re
n expresie bne
sc

bunurilor mobile i imobile, inclusiv solul, bog iile n
tur
le,
zcmintele i
lte bunuri cu poten i
l economic, disponibilit ile bneti,
Concluzie
titlurile de v
lo
re, drepturile i oblig
iile
gen ilor economici, precum
i micrile i modificrile intervenite n urm
oper
iunilor p
trimoni
le
efectu
te, cheltuielile, veniturile i rezult
tele ob inute de
ceti
.
Obiectivul fund
ment
l
l cont
bilit ii l reprezint furniz
re
de
inform
ii utile lurii deciziilor menite s
sigure o im
gine fidel

p
trimoniului, situ
iei fin
nci
re i
rezult
telor ob inute.
Din
ce
st defini ie rezult c p
trimoniul presupune existen

dou condi ii de
b
z: un titul
r de p
trimoniu (subiect de drept) pe de o p
rte, i
r pe de
lt p
rt
e bunurile i
v
lorile economice, privite c
obiecte de drepturi i oblig
ii, precum i titul
rul d
e p
trimoniu.
Structur
de
ns
mblu
p
trimoniului este urmto
re
:
PATRIMONIU
BUNURI ECONOMICE
PATRIMONIU PROPRIU
(DREPTURI)
PATRIMONIU STRIN
(OBLIGA II)
Bunurile economice se concretize
z n bunuri m
teri
le (terenuri, cldiri, util
je,
mijlo
ce de tr
nsport, stocuri de m
terii prime etc.) i nem
teri
le (proiecte, pr
ogr
me
inform
tice etc.) formnd
vere
entit ii.
Rel
iile de drepturi se refer l
f
ptul c entit
te
i procur o p
rte din
vere din
surse proprii, i
r bunurile procur
te i
p
rin de drept.
Rel
iile de oblig
ii se refer l
f
ptul c entit
te
i procur o p
rte din
vere din
surse mprumut
te, fiind oblig
t s restituie terilor contr
v
lo
re
bunurilor procur

te.
Cont
bilit
te
se ocup cu reflect
re
n expresie v
loric
p
trimoniului, e

nregistre
z circuitul elementelor p
trimoni
le n condiii concrete de timp i sp
iu,
c
lcule
z mrime

cestor elemente i reflect mic
re
p
trimoniului prin oper
iuni de
intrri i ieiri 1 .
1
Ghe. T
l
ghir, Ghe. Negoescu Cont
bilit
te
pe nelesul tuturor. Editur
All, Bucur
eti, 1998.
17Cont
bilit
te
studi
z modul de gestion
re
p
trimoniului, fund
mente
z
deciziile referito
re l
fin
n
re
i utiliz
re
elementelor p
trimoni
le, controle

z re
liz
re

deciziilor i st
bilete rspunderi privind integrit
te
i dezvolt
re
p
trimoniului.
De
semene
, cont
bilit
te
studi
z echilibrul glob
l
l p
trimoniului, prin
respect
re
ecu
iei p
trimoni
le:
BUNURI ECONOMICE = DREPTURI + OBLIGA II
cu deriv
tele s
le:
Drepturi = Bunuri economice Oblig
ii
i
Oblig
ii = Bunuri economice Drepturi
n consecin, entit
te
reprezint o unit
te p
trimoni
l,
l crei p
trimoniu po
te fi
privit sub dublu
spect:
l mijlo
celor economice (bunurile/
vere
) i
l surselor
de
procur
re

cestor mijlo
ce (c
pit
lul) proprii i strine.

Mijlo
cele economice definesc
ctivul p
trimoni
l, i
r sursele definesc p
sivul

p
trimoni
l.
n derul
re
oper
iunilor economico-fin
nci
re,
p
r i o serie de procese
economice, sub form
veniturilor i cheltuielilor, c
re
jut l
nregistr
re
creterii
s
u
diminurii p
trimoniului.
n
cest context, ecu
i
p
trimoni
l de m
i sus, devine:
AVERE = CAPITAL
182.3. Metod
cont
bilit ii
D
torit complexit ii obiectului de studiu, metod
cont
bilit ii reunete m
i multe
procedee tehnice de lucru.
Un
ns
mblu de procedee
fl
te ntr-o strns corel
ie i
Metod
intercondi ion
re c
un tot unit
r, n vedere
st
bilirii normelor i
cont
bilit ii principiilor cu c
r
cter speci
l pe c
re se fund
mente
z cont
bilit
te


i cu
jutorul cror
cercete
z st
re
i mic
re
elementelor
p
trimoni
le
le unit ilor p
trimoni
le.
O tr
stur c
r
cteristic
metodei cont
bilit ii este
cee

folosirii unor procedee
c
re s permit nregistr
re
numeric, cifric,
existen ei i micrii p
trimoniului unit
economice i soci
le n expresie v
loric. Gener
liznd, se po
te spune c metod

cont
bilit ii reprezint tot
lit
te
procedeelor interdependente, pe c
re le folosete

ce
st
n
scopul cuno
terii situ
iei p
trimoniului i
rezult
telor ob inute.
1. procedee comune tuturor tiin elor;
Procedee utiliz
te 2. procedee specifice metodei cont
bilit ii;
de cont
bilit
te 3. procedee
le metodei cont
bilit ii, comune i
ltor discipline
economice.
Procedee comune Observ
i
este f
z
ini i
l
cercetrii obiectului de studiu
l
tuturor tiin elor oricrei tiin e i este utiliz
t pentru cuno
tere
oper
iilor economic
c
re se pot exprim
v
loric i pe c
re le reflect cifric, numeric, cu

jutorul procedeelor s
le specifice.
R
ion
mentul se
plic de metod
cont
bilit ii, pentru c pe b
z
de judec i logice, pornind de l
fenomenele i procesele economice
c
re intr n obiectul su de studiu, s
jung l
concluzii noi (ex.:

ctivul este eg
l cu p
sivul, pentru c ntre mijlo
cele economice i
sursele de fin
n
re

cestor
exist o eg
lit
te perfect).
Comp
r
i
se folosete de metod
cont
bilit ii prin
ltur
re


dou s
u m
i multe fenomene i procese economice c
re se pot exprim

19v
loric, cu scopul de
st
bili
semnrile i deosebirile dintre ele, c


stfel s se tr
g o serie de concluzii. Se folosete frecvent pentru
se
comp
r
veniturile i cheltuielile pe b
z
cror
se st
bilesc rezult
tele
fin
le.
Cl
sific
re

c iune
de mpr ire, distribuire, rep
rtiz
re sistem
tic
pe cl
se s
u ntr-o
numit ordine
obiectelor n func ie de
semnrile
i deosebirile dintre ele. Asemnrile le
propie i le nc
dre
z n

cee
i cl
s, i
r deosebirile le diferen i
z i le distribuie n cl
se
diferite.
An
liz
procedeu tiin ific de cercet
re
unui ntreg,
unui fenomen
c
re se b
ze
z pe ex
min
re
fiecrui element component n p
rte.
Sintez
c
procedeu tiin ific de cercet
re
fenomenelor se b
ze
z
pe trecere
de l
p
rticul
r l
gener
l, de l
simplu l
compus pentru

se
junge l
gener
liz
re.
Procedee specifice Bil
n ul st l
b
z
dublei reprezentri
p
trimoniului n
metodei cont
bilit
te. El furnize
z inform
ii gener
le privito
re l
situ
i

cont
bilit ii economic i fin
nci
r
unei entit i, d
r reflect i l
rel
iile ei
economice cu
lte entit i, fiind complet
t de o serie de situ
ii
nex
prin c
re se explic i se det
li
z
numite l
turi
le
ctivit ii
economico-fin
nci
re
le societ ii.
Contul se deschide n cont
bilit
te
curent pentru reflect
re

fiecrui element din p
trimoniu. Cont
bilit
te
dispune de un sistem

de conturi n c
re reflect
re
oper
iilor economice rezult
te din

mic
re
elementelor p
trimoni
le
re l
b
z dubl
nregistr
re.
B
l
n
de verific
re
sigur n cont
bilit
te
dublei reprezentri i

dublei nregistrri, g
r
n i
ex
ctit ii nregistrrilor efectu
te n
conturi. D
tele b
l
n ei de verific
re st
u l
b
z
ntocmirii bil
n ului.
B
l
n
de verific
re ndeplinete
tt o func ie de control, ct i o
func ie economic, constituind punte
de legtur ntre cont i bil
n .
20Procedeele metodei Document
i
orice oper
ie economic i fin
nci
r referito
re l

cont
bilit ii comune existen
i mic
re
elementelor p
trimoni
le trebuie s fie
i
ltor discipline consemn
t n documente c
re f
c dov
d
nfptuirii lor.
economice Ev
lu
re
procedeul prin c
re d
tele cont
bilit ii sunt reprezent
te
printr-o singur unit
te de msur, crend posibilit
te
centr
lizrii
lor cu
jutorul b
l
n elor de verific
re i gener
liz
re
cu
jutorul
bil
n ului. Ev
lu
re
const n tr
nsform
re
unit ilor n
tur
le n
unit i monet
re cu
jutorul pre urilor.
C
lcul
i
este strns leg
t de ev
lu
re c
procedeu
l metodei
cont
bilit ii. Acest procedeu i gsete
plic
re
ce
m
i l
rg n
domeniul c
lcul
iei costurilor de produc ie.
Invent
riere
se folosete pentru
se cuno
te situ
i
re
l

p
trimoniului reflect
t n cont
bilit
te, i trebuie s se verifice
existen
f
ptic
tuturor elementelor s
le, n scopul descoperirii
neconcord
n elor dintre d
tele nregistr
te n conturi i re
lit
te
de
pe teren.
2.4. Principiile cont
bilit ii
Pentru
se oferi o im
gine fidel
p
trimoniului,
situ
iei fin
nci
re i
rezult

telor
ob inute de ctre entit
te, trebuie respect
te cu bun credin regulile privind ev
lu
re

p
trimoniului i celel
lte norme i principii cont
bile.
Principiul
nu sunt
dmise supr
ev
lu
re
elementelor de
ctiv i

pruden ei veniturilor, respectiv subev
lu
re
elementelor de p
siv i

cheltuielilor, innd cont de deprecierile, riscurile i pierderile
posibile gener
te de desfur
re

ctivit ii unit ii;
pruden
presupune
nticip
re
efectelor unor
c iuni i n speci
l

tr
nsferului de propriet
te cu efecte posibile
supr
exerci iului
s
u
celor p
rcurse dej
, ntruct
supr
lor nu se m
i po
te
interveni din punct de vedere cont
bil;
l
ncheiere
fiecrui exerci iu fin
nci
r se cont
bilize
z
d
toriile i pierderile prob
bile.
21Principiul
continuit
te

plicrii regulilor i normelor privind ev
lu
re
,
perm
nen ei nregistr
re

n
cont
bilit
te
i
prezent
re

elementelor
metodelor p
trimoni
le i
rezult
telor
sigurnd comp
r
bilit
te
n timp

inform
iilor cont
bile;

sigur
plic
re
pentru
cele
i elemente, stucturi, domenii de

ctivit
te,

celor
i metode de l
un exerci iu l

ltul, excluznd
schimb
re
metodelor n cursul exerci iului;
modific
re
metodelor de l
un
n l

ltul trebuie s fie
determin
t de o profund motiv
ie (modific
re
unor
cte
norm
tive, st
bilire
unor reguli gener
le de ev
lu
re noi etc.).
Principiul
presupune c unit
te
p
trimoni
l i continu n mod norm
l
continuit ii func ion
re
ntr-un viitor previzibil, fr
fi n st
re de lichid
re

ctivit ii s
u de reducere sensibil

ctivit ii;

tunci cnd func ion
re
este delimit
t n timp sunt men ion
te
d
tele de ncepere i de ncet
re

ctivit ii;

permite fix
re
respons
bilit ii entit ii

residue from
denosine triphosph
te (ATP),
which is then converted to
denosine diphosph
te (ADP). Other receptors medi
te the
remov
l of phosph
te from
phosphoryl
ted
tyrosine side ch
in by me
ns of their
phosph
t
se
ctivity (3). With one import
nt
type of receptor, lig
nd binding
ctiv
tes
gu
nyl
te cycl
se (4), which c
t
lyzes the form
tion of cyclic gu
nosine monophosph
te
(cGMP) from gu
nosine triphosph
te (GTP).
The cGMP functions
s
second messenger
nd
brings
bout
r
pid ch
nge of
ctivity of
enzymes or nonenzym
tic proteins. Remov
l or
degr
d
tion of the lig
nd reduces the concentr
tion of the second messenger
nd ends the
re
ction. (Di
gr
ms
fter Lodish et
l., 2000.) M
ny cell surf
ce receptors
ct
indirectly. When
they bind to
lig
nd they induce
series of intr
cellul
r
ctiv
tion steps. This re
ction system consists of
receptor protein,
protein (G
protein) bound to
gu
nosine residue,
nd
n
enzyme to be
ctiv
ted. Lig
nd binding
lters
the receptor protein
nd
ctiv
tes the G protein
(2). This moves to the effector, e.g.,
n enzyme
complex (3),
nd
ctiv
tes it (4). In this w
y,

second messenger is formed th
t triggers
further re
ctions in the cell, e.g., cyclic
denosine monophosph
te (cAMP) by me
ns of the
enzyme
denyl
te cycl
se (see p. 268).
References
Lodish, H. et
l.: Molecul
r Cell Biology. 4 th ed.
Scientific Americ
n Books, New York, 2000.
W
tson, J.D., et
l.: Recombin
nt DNA. 2 nd ed.
Scientific Americ
n Books, New York, 1992.
B. Hormones with immedi
te effects
on cells
Import
nt ex
mples of hormones th
t function

s lig
nds
re
mino
cid deriv
tes,
r
chidonic

cid deriv
tives,
nd m
ny peptide hormones.
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.Types of C
ell Surf
ce Receptors
P
ss
rge, Color Atl
s of Genetics 2001 Thieme
All rights reserved. Us
ge subject to terms
nd conditions of license.
267268
Genetics
nd Medicine
G Protein-coupled Receptors
The indirect tr
nsmission of sign
ls into the cell
is medi
ted by tr
nsmembr
ne proteins, which
tr
verse the cell membr
ne. A first messenger,
e.g.,
hormone like epinephrine, triggers
n intr
cellul
r re
ction by binding to
specific receptor. This le
ds to
ctiv
tion of
second messenger, which in turn initi
tes
series of re
ctions th
t result in
ch
nge of cell function.
M
ny of the genes for the different proteins involved in the indirect tr
nsmission of sign
ls

re known.
A. Stimul
tory G protein (G s )
nd

hormonereceptor complex
There
re m
ny endogenous messengers (hormones) with their own specific receptors. First
the hormone binds to the receptor (form
tion
of
hormonereceptor complex). The intr
cellul
r tr
nsmission of sign
ls is m
inly c
rried out by speci
l gu
nine-nucleotide-binding
proteins, or G proteins. By binding to gu
nosine
triphosph
te (GTP,
nucleotide composed of
gu
nine,
sug
r,
nd three phosph
te groups),
the G protein becomes
ctiv
ted
nd initi
tes
further re
ctions. G proteins consist of three
subunits: , , and . The subunit (stimul
tory
G protein, G s ) binds to the effector protein. Immedi
tely there
fter, G is in
ctiv
ted (GTP
se)
by hydrolysis of GTP to GDP (u
nosine diphosph
te). This tr
nsforms the G protein b
ck
into
n in
ctive form (G i ).
Sever
l hum
n dise
ses due to defective G protein or
defective G protein receptor
re known.
(Cl
ph
m, 1993.)
B. Four hormone cl
sses
Four princip
l cl
sses of hormones c
n be
differenti
ted: (1)
mino
cid deriv
tives such

s epinephrine
nd epinephrine deriv
tives; (2)
polypeptides such
s luc
on; (3) steroids
such
s cortisol
nd its deriv
tives;
nd (4) f
tty

cid deriv
tives such
s the prost
l
ndins.
sponsible for m
ny physioloic
l re
ctions. It
becomes in
ctiv
ted when converted into
denosine monophosph
te (AMP) by phosphodiester
se. cGMP (cyclic u
nosine monophosph
te) functions in the s
me m
nner
s
cAMP to initi
te
n intr
cellul
r re
ction.
D. G protein cycle to
ctiv
te
denyl
te
cycl
se
When
hormone binds to its specific receptor,

structur
l ch
ne occurs (1). This
ctiv
tes the
subunit of the G protein, which sep
r
tes
from the and subunits (2). The stimul
tory G
protein (G s - ) binds to the effector protein, usu
lly
denyl
te cycl
se,
nd
ctiv
tes it (3). cAMP
is then formed from ATP, while GTP is hydrolyzed to GDP
t the G- subunit. This in
ctiv
tes
the effector protein
nd the form
tion of cAMP
is termin
ted. Thus, the sin
l is of very short
dur
tion,
nd the initi
l conditions
re r
pidly
restored. Sever
l toxins exert their
ctivity by
interruptin this cycle. For ex
mple, choler

toxin inhibits in
ctiv
tion of the G s - protein so
th
t
denyl
te cycl
se rem
ins
ctiv
ted
nd
l
re
mounts of sodium
nd w
ter
re lost
throuh the intestin
l mucous membr
nes.
(Fiures
d
pted from W
tson et
l., 1992).
References
Bourne, H.R., S
nders, D.A., McCormick, F.: The
GTP
se superf
mily: conserved structure

nd molecul
r mech
nism. N
ture 349 :11
127, 1991.
Cl
ph
m, D.E.: Mut
tions in G protein-linked

receptors: novel insihts on dise
se. Cell

75 :1237 1239, 1993.
Linder, M.E., Gilm
n, A.G.: G-Proteins, Sci. Am.
36 43, 1992.
W
tson, J.D., et
l.: Recombin
nt DNA. 2 nd ed.
W.H. Freem
n, Scientific Americ
n Books,
New York, 1992.
C. Form
tion
nd hydrolysis of cAMP
The key re
ction is the form
tion of cyclic
denosine monophosph
te (cAMP) from
denosine
triphosph
te (ATP) by me
ns of
denyl
te cycl
se. Intr
cellul
r cyclic AMP tr
nsmits the
ctiv
tion initi
ted by the hormonereceptor
complex without
molecule h
vin p
ssed
throuh the pl
sm
membr
ne. cAMP is reP
ss
re, Color Atl
s of Genetics 2001 Thieme
All rihts reserved. Us
e subject to terms
nd conditions of license.269
G Protein-coupled Receptors
Hormone
H
Receptor
Lipid bil
yer
H
Receptor
Receptor
"
G protein
in
ctive
"
G protein
ctive (G s )
GTP
Bindin to G protein
!
#
!
#
Hormone-receptor complex
H
Receptor
"
!
In
ctiv
tion
of G ! (GTP
se)
Effect
on
effector
protein
!
#
Activ
tion of G !
GDP
A. Stimul
tory G protein (Gs)
nd hormone-receptor complex
1 Norepinephrine
Phosphodiester
se
Adenyl
te cycl
se
(
mino
cid deriv
tive)
Adenine
2 Gluc
on
Adenine
Adenine

(polypeptide)
P
3 Cortisol
(steroid)
4 Prost
l
ndin A 2
P
P
Ribose
Adenosine
triphosph
te (ATP)
(f
tty
cid deriv
tive)
B. Four hormone types
#
G protein
Cyclic
denosine
monophosph
te (cAMP)
Ribose
Adenosine
monophosph
te
(AMP)
H
Receptor
"
P
Ribose
C. Form
tion
nd hydrolysis of cAMP
Effector protein
e..,
denyl
te cycl
se
H
P
Receptor
"
!
#
!
G s
GTP
1. G protein binds to hormone-receptor
1. complex
2. G protein
ctiv
ted
H
Receptor
Receptor
"
#
"
#
!
GDP
!
ATP
cAMP
G s in
ctiv
ted by GTP
se
4. G protein in
ctiv
ted
3. Effector protein
ctiv
ted
D. G protein cycle to
ctiv
te
denyl
te cycl
se
P
ss
re, Color Atl
s of Genetics 2001 Thieme
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e subject to terms
nd conditions of license.
Physioloic
l

effect270
Genetics
nd Medicine
Function
l sin
ls between cells
re received by
tr
nsmembr
ne proteins
s sin
l tr
nsmitters.
Durin evolution, rel
tively simple precursor
enes for such proteins 
ve rise to multiple
structur
lly
nd function
lly rel
ted enes.
Their correspondin proteins serve to tr
nsmit
ions (sodium, pot
ssium, c
lcium, chloride,
nd
others),
s neurotr
nsmitters,
nd for perception of liht
nd odors, etc. Clonin of these
enes h
s yielded insiht into the v
riety of
functions of tr
nsmembr
ne sin
l tr
nsmitters. Their ener
l structure c
n be tr
ced b
ck
to
n evolution
rily conserved
ncestr
l
molecule. follow. An especi
lly common structur
l motif
is
tr
nsmembr
ne protein cont
inin seven
helices within the pl
sm
membr
ne. The

mino end is extr
cellul
r; the c
rboxy end is
intr
cellul
r. Different olios
cch
ride side
ch
ins
re usu
lly bound to the extr
cellul
r
dom
ins. The intr
cellul
r dom
ins h
ve bindin sites for other molecules involved in sin
l
tr
nsmission. The seven-helix motif is the
ch
r
cteristic structure of G protein-bindin receptors (p. 268). As the G proteins themselves,
these receptors
nd their enes form
l
re
f
mily with
lon evolution
ry history. In
ye
st, they serve to discern the pheromones of
the m
tin types (p. 186); in hiher or
nisms
they
re the b
sis for tr
nsmittin sin
ls of vision, smell,
nd t
ste (p. 278 286). (Fiure redr
wn from Stryer, 1995).
A. Tr
nsmembr
ne structure of
volt
e-
ted ion ch
nnels C. A receptor with two tr
nsmembr
ne protein ch
ins,
nd
The direct flow of ions across the cell memrane
is regulated y ion channels. The transmemrane proteins, composed of several domains,
are arranged so that they form pores that can e
opened and closed. The simplest model is the
potassium channel (1). This memrane-ound
polypeptide contains six transmemrane
domains. The amino and the caroxy ends of
the protein lie within the cell. Changes in cell
memrane potential or voltage cause the channel to open (or close) in order to initiate (or terminate) a rief flow of ions. Domain 4, which is
composed of polar amino acids, is crucial for the
flow of ions. Sodium and calcium ion channels
consist of four suunits (2) of similar structure,
each resemling a potassium channel. The similarity is due to the common evolutionary origin
of their genes. The four suunits of the sodium
channel (3) are positioned to form a very narrow porelike passage, much narrower than a
potassium channel, through the plasma memrane. Ion transport is rought aout y memrane depolarization (3 and 4). (Figure after
Watson et al., 1992.) The receptor for -
minobutyric
cid (GABA)

utilizes two tr
nsmembr
ne protein subunits,

nd . Both the amino and the caroxy ends are
extracellular. The two chains are coded for y
different genes. Several oligosaccharide side
chains are present on the extracellular side. The
chain contains a phosphorylation site for
cAMP-dependent protein kinase.
(Figures adapted from Watson et al., 1992).
Transmemrane Signal
Transmitters
References
Saatini, D.D., Adesnik, M.B.: The iogenesis of
memranes and organelles. pp. 459 553,
In: Scriver, C.R., et al., eds., The Metaolic
and Molecular Bases of Inherited Disease.
7 th ed. McGraw-Hill, New York, 1995.
Stryer, L.: Biochemistry, 4 th ed. Freeman Pul.,
San Francisco, 1995.
Watson, J.D. et al.: Recominant DNA, 2 nd ed.,
Scientific American Books, New York, 1992.
B. Seven-helix structure of
transmemrane signal transmitters
Indirect transmission of signals is more
frequent than the direct transport of ions or ligand-gated impulse transmission. Here, the
transmemrane protein is involved only in the
first step of signal transmission. Further steps
Passarge, Color Atlas of Genetics 2001 Thieme
All rights reserved. Usage suject to terms and conditions of license.Transmemr
ane Signal Transmitters
Passarge, Color Atlas of Genetics 2001 Thieme
All rights reserved. Usage suject to terms and conditions of license.
271272
Genetics and Medicine
Receptors of Neurotransmitters
Impulses are relayed etween nerve cells or etween nerve and muscle cells y various transmitter molecules (neurotransmitters). Their effects are further relayed y receptors in the cell
memrane. Receptors can e differentiated according to their structure, which in turn determines their specificity.
A. Acetylcholine as a neurotransmitter
Cholinergic synapses convey the nerve impulse
from one nerve cell to another or from a nerve
cell to a muscle cell (motor endplate). Acetylcholine leads to postsynaptic depolarization
through the release of potassium ions (K + ) and
the uptake of sodium ions (Na + ). This process is
regulated y an acetylcholine receptor.
B. Acetylcholine receptors
The acetylcholine receptors are of two genetically and functionally different types. Pharmacologically they can e differentiated according to the effects of nicotine and muscarine.
The nicotine-sensitive acetylcholine receptor is
an ion channel for potassium and sodium. It
consists of five suunits: two , one , one ,

nd one (1). Acetylcholine bins as a ligan to
the two subunits. E
ch subunit consists of

four tr
nsmembr
ne dom
ins (2). E
ch subunit

is encoded by its own ene (3). These enes
h
ve simil
r structures
nd nucleotide b
se
sequences. The li
nd-
ted ion ch
nnel is
n
ex
mple of direct tr
nsport without
n intermedi
te c
rrier. A mut
tion in the second tr
nsmembr
ne reion h
s been shown to ch
ne
the ion selectivity from c
tions to
nions (G
lzi,
1992.)
The musc
rine-sensitive type of
cetylcholine
receptor is
protein th
t conti
ns seven tr
nsmembr
ne subunits (4). Since e
ch exists in the
form of
n helix, it is referred to
s
sevenhelix tr
nsmembr
ne protein (p. 270). The

mino end (NH 2 ) lies extr
cellul
rly; the c
rboxy end (COOH), intr
cellul
rly. The tr
nsmembr
ne dom
ins
re connected by intr
cellul
r
nd extr
cellul
r polypeptide loops (4).
Different dom
ins of the whole protein
re distinuished (5)
ccordin to loc
tion
nd the
rel
tive proportion of hydrophilic
nd hydrophobic
mino
cids. The
mino end
nd the c
rboxy end e
ch form
dom
in just like the intr
cellul
r (
c),
nd extr
cellul
r portions (df).
The tr
nsmembr
ne dom
ins loc
ted within
the pl
sm
membr
ne (1 7) consist for the
most p
rt of hydrophobic
mino
cids. The
structure of the ene product corresponds to
the ener
l structure of the ene (6). The different dom
ins
re coded for by individu
l exons.
The DNA nucleotide sequences within function
lly simil
r dom
ins
re simil
r.
The seven-helix tr
nsmembr
ne motif occurs
in m
ny receptors. The ener
l structures of the
enes
nd of the ene products
re very simil
r,
but they differ in their specificity of bindin to
other function
lly relev
nt molecules (G proteins). They pl
y
role not only
s neurotr
nsmitters but
lso in the tr
nsmission of
liht, odors,
nd t
ste. (Fiures b
sed on W
tson et
l., 1992.)
References
G
lzi, G.L.: Mut
tions in the ch
nnel dom
in of

neuron
l nicotinic receptor convert ion
selectivity from c
tionic to
nionic. N
ture
359 :500 505, 1992.
W
tson, J.D., Gilm
n, M., Witkowski, J., Zoller,
M.: Recombin
nt DNA. 2 nd ed. W.H.
Freem
n, Scientific Americ
n Books, New
York, 1992.
P
ss
re, Color Atl
s of Genetics 2001 Thieme
All rihts reserved. Us
e subject to terms
nd conditions of license.273
Receptors of Neurotr
nsmitters
Presyn
ptic
Acetylcholine
+
+
K+
+ + - ++ +

+ + +
- - - - - K + Hih
N
+ Low
N
+
Pol
rized
Postsyn
ptic
Depol
rized
Cholineric syn
pse (nerve/nerve or nerve/muscle)
A. Acetylcholine
s neurotr
nsmitter
Two types of
cetylcholine receptors
Seven-helix tr
nsmembr
ne protein
bound to G proteins
(musc
rine-sensitive)
C
tion-specific ch
nnel
in muscle of vertebr
tes
(nicotine-sensitive)
1. Acetylcholine
1. binds to
1. !-subunits
1. (li
nd
1. bindin)
C
tions
(K +, N
+ )
$
"
!
extr
cellul
r
d
NH2
3
4.
1
#
2
e
5
4
f
6
7
intr
cellulr


5.
2. E
ch subunit
2. h
s four
2. tr
nsmembr
ne
2. dom
ins
A
COOH
b
c
1
2 d 3 b 4 e 5 c 6 f 7
A
B

- c
d - f
1 - 7

B
Amino end
C
rboxy end
intr
cellul
r dom
ins
extr
cellul
r dom
ins
tr
nsmembr
ne dom
ins (hydrophobic)
6. Gene structure (di
r
m)
3. One ene for e
ch subunit:
2 for
1 for
1 for
1 for
!-subunits
"-subunit
#-subunit
$-subunit
Exons
5'
A
Introns
1
2 d 3 b 4 e 5 c 6 f 7
The different dom
ins
re encoded
by individu
l exons
B. Acetylcholine receptor
P
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re, Color Atl
s of Genetics 2001 Thieme
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e subject to terms
nd conditions of license.
3'
B274
Genetics
nd Medicine
Genetic Defects in Ion Ch
nnels
More th
n 20 different disorders due to defective ion ch
nnel proteins resultin from ene
mut
tions
re known. Such disorders include
cystic fibrosis (see p. 276), the lon-QT syndrome,
speci
l type of de
fness, heredit
ry
hypertension (Liddle syndrome), f
mili
l persist
nt hyperinsulinemic hypolycemi
of inf
ncy, some heredit
ry muscle dise
ses,
nd
m
lin
nt hyperthermi
(see p. 372),
mon
other disorders.
A. Lon-QT syndrome,
enetic
c
rdi
c
rrhythmi

Conenit
l lon-QT syndrome is ch
r
cterized
by
proloned QT interv
l in the electroc
rdior
m (more th
n 460 ms, corrected for he
rt
r
te), sudden
tt
cks of missed he
rt be
ts
(syncopes) or series of r
pid he
rt be
ts (tors
des de pointes),
nd
n incre
sed risk for sudden de
th from ventricul
r fibrill
tion in
children
nd youn
dults.
B. Different molecul
r types of
lon-QT syndrome
Prolon
tion of the QT interv
l in the electroc
rdior
m results from
n incre
se in the dur
tion of the c
rdi
c
ction potenti
l (1). The
norm
l potenti
l l
sts
bout 300 ms (ph
ses 1

nd 2). The restin membr
ne potenti
l (ph
se
3) is re
ched by proressive in
ctiv
tion of c
lcium currents
nd incre
sin depletion of
pot
ssium currents, which repol
rize the cell.

In ph
se 0 the cell is quickly depol
rized by
ctiv
ted sodium currents followin
n excit
tory
stimulus.
LQT1
ccounts for
bout h
lf of the p
tients
with lon-QT syndrome. The ene for LQT2 encodes
1195-
mino-
cid tr
nsmembr
ne protein responsible for the other m
jor pot
ssium
ch
nnel th
t p
rticip
tes in ph
se 3 repol
riz
tion (HERG st
nds for (hum
n-ether-r-o-orel
ted ene,
Drosophil
homoloue). LQT3,

sodium ch
nnel protein, consists of four subunits, e
ch cont
inin six tr
nsmembr
ne
dom
ins
nd
number of phosph
te-bindin
sites. Homozyosity for LQT1 (KVLQT1 ene
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e subject to terms
nd conditions of license.227
Genomic Imprintin
dies very
e
rly
2
Androenetic
1
norm
l
development
Extr
embryonic
tissues
3
Diploid zyote
Fetus
bsent
or stunted
Preimpl
nt
tion
f
ilure in most
Norm
l
Fetus norm
l
Preimpl
nt
tion
dies
l
ter
4
Fetus norm
l
until 40 somite
st
e
Preimpl
nt
tion norm
l,
extr
-embryonic
tissues underdeveloped
Gynoenetic
A. The import
nce of two different p
rent
l enomes
Two p
tern
l enomes
1. Hyd
tidiform mole
Two m
tern
l enomes
2. Hyd
tidiform mole
3. Ov
ri
n ter
tom

4. Triploidy 69, XXX
B. Hum
n embryonic development depends on presence of
m
tern
l
nd
p
tern
l
enome
1.
P
Som
tic cells
XX
nd XY
M
le

P
2.
P P
tern
l
M
In
ctive Active
Active In
ctive
P
M M
tern
l
Fem
le
M
M
Imprint
er
sed
Primordi
l
erm cells
P
M
3.
Imprint
reset
G
metes
P
M
4.
Zyote
C. Genomic imprintin is est
blished in e
rly embryonic development
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re, Color Atl
s of Genetics 2001 Thieme
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e subject to terms
nd conditions of license.
Imprint
est
blished228
Fund
ment
ls
X-Chromosome In
ctiv
tion
Durin e
rly embryonic development of m
mm
li
n fem
les, one of the two X chromosomes
becomes in
ctiv
ted. The in
ctiv
tion is induced by
ene (XIST, X-in
ctiv
tion-specific
tr
nscript) on the proxim
l lon
rm of the X
chromosome, tr
nscribed from the in
ctive X.
X in
ctiv
tion is
mech
nism to b
l
nce Xchromosom
l ene expression between fem
le

nd m
le cells.
A. X chrom
tin
In 1949, B
rr und Bertr
m observed
st
in
ble

ppend
e in the nucleus of nerve cells of
fem
le (1
nd 3) but not of m
le c
ts (2). The

uthors n
med this structure
s sex chrom
tin. Simil
r structures were found drumsticks in peripher
l blood leukocytes (4)
nd
sm
ll peripher
l bodies in the nuclei of fibrobl
sts
nd or
l mucos
cells (5) in hum
ns. E
ch
of these structures represents one of the two X
chromosomes
nd
re referred to
s X chrom
tin. (Fiures 1 3 from B
rr
nd Bertr
m,
1949).
B. Scheme of X in
ctiv
tion
R
ndom in
ctiv
tion of most of the enes of one
of the two X chromosomes in fem
le cells
occurs e
rly in embryoenesis,
t
bout d
y 21
in hum
ns. In
iven cell, it involves the X chromosome of either m
tern
l or p
tern
l oriin.

The in
ctiv
tion p
ttern is norm
lly irreversible

nd st
bly tr
nsmitted to
ll d
uhter cells. The
expected distribution is usu
lly 1 : 1. In r
re inst
nces this m
y be skewed tow
rd
preferenti
l type of in
ctiv
tion. In extreme c
ses this
m
y result in clinic
l m
nifest
tion in
heterozyous fem
le if the m
jority of cells cont
in
the mut
tion on the
ctive X chromosome.
C. Mos
ic p
ttern of expression
Fem
le som
tic tissues show
mos
iclike distribution of cells expressin just one of the two

lleles. In mice, X-chromosom
l co
t-color mut
nts show
mos
ic of liht-
nd d
rk-colored
co
t p
tches (1,
fter Thompson, 1965). In
hum
ns,
simil
r distribution of norm
l
nd

bsent swe
t pores is seen in fem
le heterozyotes for hypohidrotic ectoderm
l dyspl
si
.
The swe
t pores of hemizyotes
re
bsent
(hypohidrosis). Finerprints of fem
le heterozyotes show
re
s with norm
l swe
t pores
(bl
ck points)
nd
re
s with
bsent swe
t pores
(2

nd 2b, from P
ss
re
nd Fries, 1973). In
cell cultures from fem
le heterozyotes for
X-chromosom
l HGPRT deficiency (hypox
nthineu
ninephosphoribosyltr
nsfer
se)
the colonies
re either HGPRT or HGPRT + (3).
(From Mieon, 1971, with kind permission of the

uthor
nd publisher).
D. Exceptions from X-in
ctiv
tion
Tr
nscription
l silencin of X-chromosom
l
enes in m
mm
li
n fem
le cells is not
complete. Some enes esc
pe in
ctiv
tion
nd

re expressed from both the
ctive
nd the in
ctive X chromosome. The m
jority of these enes
h
ve
homoloue on the Y chromosome,
reflectin
common evolution
ry oriin. P
nel
D shows enes th
t
re expressed on the in
ctive hum
n X chromosome. Most
re loc
ted
t
the ends of the X chromosomes. (Fiure

d
pted from Brown et
l., 1997).
E. X-in
ctiv
tion profile
An
n
lysis of 224 X-linked tr
nscripts showed
th
t 34 esc
pe in
ctiv
tion (C
rrel et
l., 1999).
Of these, 31 m
p to the short
rm of the X chromosome. The expressed enes
re open circles,
the in
ctiv
ted enes filled circles. Sever
l
enes in the pseudo
utosom
l rei
observ
tions
re correct, the future of life
nd the universe will be f
r ble
ke
r.
In the l
st four ye
rs
stronomers h
ve reported evidence th
t the exp
nsion of
the
universe is not just continuin but is speedin up, under the influence of
mys
terious
"d
rk enery,"
n
ntir
vity th
t seems to be embedded in sp
ce itself. If th
t
is true
nd
the universe oes on
cceler
tin,
stronomers s
y, r
ther th
n co
stin ently
into the
niht, dist
nt 
l
xies will eventu
lly be movin
p
rt so quickly th
t they c
n
not
communic
te with one
nother. In effect, it would be like livin in the middle o

f
bl
ck
hole th
t kept ettin emptier
nd colder.
In such
universe, some physicists s
y, the usu
l methods of formul
tin physic
s m
y
not
ll
pply. Inste
d of new worlds comin into view, old ones would const
ntly
be
dis
ppe
rin over the horizon, lost from view forever.
Cosmoloic
l knowlede would be fr
mented, with different observers doomed to
seein different pieces of the puzzle
nd no sinle observer
ble to know the f

te of the
whole universe or
rrive
t
theory of physics th
t w
s more th
n
pproxim
te.
37"There would be
lot of thins
bout the universe th
t we simply couldn't pre
dict," s
id
Dr. Thom
s B
nks,
physicist
t the University of C
liforni

t S
nt
Cruz.
And perh
ps most import
nt, st
rved fin
lly of the enery even to complete
tho
uht or

comput
tion, the dom
in of life
nd intellience would not exp
nd, but constri
ct
nd
eventu
lly v
nish like
dwindlin echo into the silence of eternity. "I find th
e f
te of

universe th
t is
cceler
tin forever not very
ppe
lin," s
id Dr. Edw
rd Witte
n,

theorist
t the Institute for Adv
nced Study.
Th
t is
n underst
tement, in the view of Dr. L
wrence M. Kr
uss,
n
strophysic
ist
t
C
se Western Reserve University in Clevel
nd, who
lon with his colle
ue Dr. G
lenn
D. St
rkm
n h
s recently tried to limn the possibilities of the f
r future. An

cceler
tin
universe "would be the worst possible universe, both for the qu
lity
nd qu
ntit
y of life,"
Dr. Kr
uss s
id,
ddin: "All our knowlede, civiliz
tion
nd culture
re destin
ed to be
forotten. There's no lon-term future."
Einstein's L
st L
uh
Until
bout four ye
rs
o,
n overwhelmin preponder
nce of
stronomers subscri
bed to
the view th
t the cosmic exp
nsion w
s prob
bly slowin down bec
use of the coll
ective
r
vity of the 
l
xies
nd everythin else in the universe, the w
y
h
ndful o
f stones
tossed in the
ir r
du
lly slow their
scent. The only question w
s whether the
universe
h
d enouh r
vit
tion
l oomph to stop exp
ndin
nd brin itself b
ck toether
in
"bi
crunch," or whether the 
l
xies would s
il ever more slowly outw
rd forever.
It w
s to me
sure th
t r
te of slowin of this outw
rd fliht,
nd thus find the
lon-souht

nd elusive
nswer to the cosmic question, th
t two te
ms of
stronomers st
rted
competin projects in the 1990's usin dist
nt explodin st
rs, supernov
s,
s c
osmic
be
cons.
In 1998 the two te
ms
nnounced th
t inste
d of the expected slowin, the 
l
xi
es

ctu
lly seem to h
ve speeded up over the l
st five or six billion ye
rs,
s if
some "d
rk
enery" w
s pushin them outw
rd.
"It's definitely the str
nest experiment
l findin since I've been in physics,"
Dr. Witten
s
id. "People find it difficult to
ccept. I've stopped expectin th
t the findi

n will be
proved wron, but it's
n extremely uncomfort
ble result."
To
stronomers this d
rk enery be
rs
h
untin resembl
nce to
n ide
th
t Alb
ert
Einstein h
d b
ck in 1917
nd then
b
ndoned, l
ter c
llin it his biest blund
er. In th
t
ye
r he inserted
m
them
tic
l fude f
ctor th
t c
me to be known
s the cosmol
oic
l
const
nt into his equ
tions of ener
l rel
tivity in order to st
bilize the univ
erse

inst
coll
pse; Einstein's const
nt
cted
s
kind of cosmic repulsion to b
l
nce the
r
vit
tion
l pull of the 
l
xies on one
nother.
38Einstein 
ve up the cosmoloic
l const
nt
fter the Americ
n
stronomer Edwin
Hubble
discovered th
t the universe w
s exp
ndin
nd thus did not need st
bilizin. Bu
t his
fude f
ctor refused to die. It 
ined
new identity with the
dvent of qu
ntum
mech
nics, the biz
rre-soundin rules th
t overn the sub
tomic re
lm. Accordin
to
those rules, empty sp
ce is not empty, but r
ther fo
min with enery. Inserted
into
Einstein's equ
tions, this enery would
ct like
cosmoloic
l const
nt,
nd tr
y to blow
the universe
p
rt.
Accordin to
stronomers the recently discovered d
rk enery now
ccounts for
b
out
two-thirds of the m
ss of the universe. But is this Einstein's old fude f
ctor,
the
cosmoloic
l const
nt, come home to roost -- in which c
se the universe will
cc
eler
te
etern
lly? Or is the presumed
cceler
tion only tempor
ry, driven by one of the
m
ny
mysterious force fields, dubbed quintessence,
llowed by v
rious theories of hi
h enery
physics?
Or is the
cceler
tion even re
l?
"It's import
nt to find out if the cosmoloic
l const
nt is re
lly const
nt," s

id Dr. Witten.
Bec
use the repulsive force resides in sp
ce itself,
s the universe rows, the
push from
d
rk enery rows
s well. "If d
rk enery is the cosmoloic
l const
nt then it
is

property of the v
cuum th
t will
lw
ys be with us, ettin more powerful
s the
universe
ets bier
nd the universe will exp
nd forever," expl
ined Dr. Ad
m Riess of t
he Sp
ce
Telescope Science Institute in B
ltimore. But if the d
rk enery is some form of
quintessence, "then there m
y be more such fields which
rise in the future, pos
sibly of
the opposite sin,
nd then
ll bets
re off for the future of the universe."
Dr. Kr
uss s
id, "The ood news is th
t we c
n't prove th
t this is the worst of

ll
possible universes."
The Lon Goodbye
It miht seem str
ne or presumptuous for
stronomers to try to describe events

ll the
w
y to the end of time when physicists
re still ropin for
"theory of everyt
hin." But
to Dr. Kr
uss, this is testimony to the power of ordin
ry physics. "We c
n still
put

ultim
te limits on thins without even knowin the ultim
te theory," he s
id. "W
e c
n put
limits on thins b
sed on ordin
ry physics."
Dr. Dyson s
id his venture into esch
toloy w
s inspired p
rtly by
1977 p
per
on the
future of
n ever exp
ndin universe by Dr. J. N. Isl
m, now
t the University o
f
Chitt
on in B
nl
desh, in The Qu
rterly Journ
l of the Roy
l Astronomic
l Soc
iety.
Dr. Dyson w
s
lso motiv
ted, he wrote in his p
per, to provide
counterpoint t
o

f
mously dour st
tement by Dr. Steven Weinber, who wrote in his book "The First
Three Minutes," "The more the universe seems comprehensible, the more it
lso se
ems
pointless."
39Dr. Dyson wrote, "If Weinber is spe
kin for the 20th century, I prefer the 1
8th."
If the present trend of
cceler
tion continues this is the forec
st:
In
bout two billion ye
rs E
rth will become uninh
bit
ble
s
r
du
lly w
rmin
 Sun
produces
run
w
y reenhouse effect. In five billion ye
rs the Sun will swell u
p
nd die,
burnin the E
rth to
crisp in the process. At
bout the s
me time the Milky W

y will
collide with its twin the Andromed

l
xy, now
bout two million liht-ye
rs
w

y
nd
closin f
st, spewin st
rs, 
s
nd pl
nets
cross inter
l
ctic sp
ce.
Any civiliz
tion th
t m
n
ed to survive these events would f
ce
future of inc
re
sin
inor
nce
nd d
rkness
s the
cceler
tin cosmic exp
nsion rushes most of the u
niverse

w
y from us. "Our
bility to know
bout the universe will decre
se with time,"
s
id Dr.
Kr
uss. "The loner you w
it, the less you see, the opposite of wh
t we
lw
ys t
houht."
As he expl
ins it, the dis
ppe
r
nce of the universe is
r
du
l process. The f

ster


l
xy flies
w
y from us, the dimmer
nd dimmer it will
ppe
r,
s its liht is
"redshifted" to lower frequencies
nd eneries, the w
y
police siren sounds lo
wer when
it is recedin. When it re
ches the speed of liht, the 
l
xy will
ppe
r to "f
reeze," like

d
ncer c
uht in mid
ir in
photor
ph, in
ccord
nce to Einstein's theory of r
el
tivity,

nd we will never see it et older, s
id Dr. Abr
h
m Loeb,
n
stronomer
t H
rv

rd.
R
ther it will simply seem dimmer. The f
rther
w
y
n object is in the sky, he
s
id, the
youner it will
ppe
r
s it f
des out of siht. "There is
finite
mount of in
form
tion we
c
n collect from the universe," Dr. Loeb s
id. About 150 billion ye
rs from now

lmost

ll of the 
l
xies in the universe will be recedin f
st enouh to be invisible
from the
Milky W
y. The exceptions will be 
l
xies th
t
re r
vit
tion
lly bound to the
cloud of

l
xies, known
s the Loc
l Group, to which the Milky W
y belons. Within this
cloud,
life would look much the s
me
t first. There would be 
l
xies in the sky. "Whe
n you

look
t the niht the st
rs will still be there," s
id Dr. Kr
uss. "To the
stro
nomer who
w
nts to see beyond, the sky will be s
dly empty. Lovers won't be disturbed -- s
cientists
will be."
But
bout 100 trillion ye
rs from now, when the interstell
r 
s
nd dust from w
hich new
st
rs condense is fin
lly used up, new st
rs will ce
se to be born. From th
t ti
me on, the
sky will row d
rker
nd d
rker. The 
l
xies themselves,
stronomers s
y, will
coll
pse
in bl
ck holes within
bout 1030 ye
rs.
But even
bl
ck hole is not forever,
s Dr. Stephen H
wkin, the C
mbride Univ
ersity
physicist
nd best-sellin
uthor, showed in p
th-bre
kin c
lcul
tions b
ck in
1973.
Applyin the principles of qu
ntum mech
nics to these dre
d-soundin objects, Dr
.
H
wkin discovered th
t
bl
ck hole's surf
ce, its so-c
lled event horizon, wou
ld
fluctu
te
nd exude enery in the form of r
ndom bursts of p
rticles
nd r
di
ti
on,
rowin hotter
nd hotter until the bl
ck hole eventu
lly exploded
nd v
nished.
Bl
ck holes the m
ss of the sun would t
ke 1064 ye
rs to explode. For bl
ck hole
s the
m
ss of

l
xy those fireworks would liht up sp
ce-time 1098 ye
rs from now.
40A
inst the F
ll of Niht
Will there be
nythin or
nyone
round to see these qu
ntum fireworks?
Dr. Dyson
rued in his 1979 p
per th
t life
nd intellience could survive the
desert of
d
rkness
nd cold in
universe th
t w
s exp
ndin infinitely but ever more slow
ly by

doptin ever slower
nd cooler forms of existence. Intellience, could reside,
for
ex
mple, in the p
ttern of electric
lly ch
red dust r
ins in
n interstell
r c
loud,

situ
tion described in the 1957 science fiction novel "The Bl
ck Cloud," by the
British

stronomer Sir Fred Hoyle, who died in Auust.
As
n or
nism like the bl
ck cloud cooled, he
rued, it would think more slowl
y, but it
would
lw
ys met
bolize enery even more slowly, so its
ppetite would
lw
ys be
less
th
n its output. In f
ct, Dr. Dyson concluded, by m
kin the
mount of enery ex
pended
per thouht sm
ller
nd sm
ller the cloud could h
ve
n infinite number of thou
hts
while consumin only
finite
mount of enery.
But there w
s
hitch. Even just thinkin requires enery
nd ener
tes he
t, wh
ich is
why computers h
ve f
ns. Dr. Dyson suested th
t cre
tures would h
ve to stop
thinkin
nd hibern
te periodic
lly to r
di
te
w
y their he
t.
In
n
cceler
tin universe, however, there is
n
ddition
l source of he
t th
t
c
nnot be
otten rid of. The s
me c
lcul
tions th
t predict bl
ck holes should explode
ls
o predict
th
t in
n
cceler
tin universe sp
ce should be filled with so-c
lled H
wkin r

di
tion.
In effect, the horizon -- the f
rthest dist
nce we c
n see -- looks m
them
tic
l
ly like the

surf
ce of
bl
ck hole. The
mount of this r
di
tion is expected to be incredib

ly sm
ll -correspondin to
fr
ction of
billionth of
billionth of
billionth of
de
ree
bove

bsolute zero, but th
t is enouh to doom sentient life.
"The H
wkin r
di
tion kills us bec
use it ives
minimum temper
ture below whi
ch
you c
nnot cool
nythin," s
id Dr. Kr
uss. Once
n or
nism cools to th
t tempe
r
ture,
he expl
ined, it would dissip
te enery
t some fixed r
te. "Since there is
fi
nite tot
l
enery, this me
ns
finite lifetime."
Infinity on Tri
l
Althouh Dr. Dyson
rees with this loomy view of life in
n
cceler
tin unive
rse, he

nd Dr. Kr
uss
nd Dr. St
rkm
n
re still
ruin
bout whether life is
lso doo
med in

universe th
t is not
cceler
tin, but just exp
ndin
nd ettin slower
nd col
der.
Qu
ntum theory, the C
se Western
uthors point out, limits how finely the enery
for
new thouhts c
n be sh
ved. The theory decrees th
t enery is emitted
nd
bsorb
ed in
tiny indivisible lumps c
lled "qu
nt
." Any comput
tion must spend
t le
st this
much
enery out of
limited supply. E
ch new thouht is
step down
n enery l
dder
with

41finite number of steps. "So you c
n only h
ve
finite number of thouhts," s

id Dr.
Kr
uss.
"If you w
nt to st
re
t your n
vel
nd not think
ny new thouhts, you won't di
ssip
te
enery, " he expl
ined. But th
t would be
borin w
y to spend eternity. If lif
e is to
involve more th
n the etern
l reshufflin of the s
me d
t
, he
nd Dr. St
rkm
n
s
y, it
c
nnot be etern
l.
Dr. Dyson, however, s
ys this
rument
pplies only to so-c
lled diit
l life, i
n which
there is
fixed number of qu
ntum st
tes. Cre
tures like the bl
ck cloud, which
could
row
lon with the universe, he s
id, would h
ve
n incre
sin number of qu
ntu
m
st
tes,
nd so there would
lw
ys be more runs of the l
dder to step down. So t
he
bottom need never be re
ched
nd life
nd thouht could o on indefinitely.
But nobody knows whether such
life form c
n exist, s
id Dr. Kr
uss.
Comp
red with the siht of the World Tr
de Center towers coll
psin or the plih
t of

sick child, this future extinction m
y seem
remote concern. Dr. All
n S
nd
e,

n

stronomer
t C
rneie Observ
tories in P
s
den
, C
lif., who h
s spent his life
investi
tin the exp
nsion
nd f
te of the universe, s
id: "Life on this e
rth
is oin to
v
nish in 4.5 billion ye
rs. I wouldn't et hun up on the f
ct th
t the lihts

re
ll oin
out in 30 billion ye
rs."
Dr. Dyson s
id he w
s still
n optimist. It is too soon to st
rt p
nickin, he c
ounseled in

n e-m
il mess
e. The observ
tions could be wron.

"At present
ll possibilities
re open," he wrote. "The recent observ
tions
re
import
nt,
not bec
use they
nswer the bi questions
bout the history of the universe, but
bec
use
they ive us new tools with which to explore the history."
Even in
n
cceler
tin universe, Dr. Dyson s
id, hum
ns or their descend
nts mi
ht one
d
y be
ble to re
rr
ne the 
l
xies
nd s
ve more of them from dis
ppe
rin. A
nother
limmer of hope comes from the de
dly
nd chillin H
wkin r
di
tion itself, s
i
d Dr.
R
ph
el Bousso, from the Institute of Theoretic
l Physics
t the University of C

liforni


t S
nt
B
rb
r
. Since th
t r
di
tion is produced by unpredict
ble qu
ntum fluc
tu
tions,
he pointed out, if you w
it lon enouh
nythin c
n
ppe
r in it, even
new un
iverse.
"Sooner or l
ter one of those qu
ntum fluctu
tions will look like
Bi B
n," h
e s
id.
In th
t c
se there is the possibility of
future, if not for us,
t le
st for s
omethin or
somebody. In the fullness of time,
fter
ll, physics te
ches th
t the improb
bl
e
nd even
the seeminly impossible c
n become the inevit
ble. N
ture is not done with us y
et, nor,

s Dr. Dyson indic
tes,
re we necess
rily done with n
ture.
We
ll die,
nd it is up to us to decide who
nd wh
t to love, but,
s Dr. Weinb
er
pointed out in
recent
rticle in The New York Review of Books, there is
cert

in
nobility in th
t prospect.
42"Thouh
w
re th
t there is nothin in the universe th
t suests
ny purpose
for
ure brins
bout
n incre
se of the thickness
nd
shortenin of the surf
ce, while, on the
other h
nd, tension le
ds to splittin of the continent
l blocks. The individu
l st
es of perceived
s
mount
in form
tion comprised continu
l processes
of splittin
nd compression, whereby the oriin
l
Si
lic crust (for which Weener
ssumed
thickness
of
bout 3035 km) r
du
lly decre
sed in surf
ce

re
, split into sep
r
te pieces,
nd incre
sed in thickness. Alon with the movement of the continent
l
blocks,
hypothesized univers
l oce
n (P
nth
l
ss
)
began to ivie into a shallow sea an a eep sea.
Volcanism, for Wegener, was mainly relate to the
continental fronts. Areas where tension prevaile,
such as the Atlantic Ocean, an also opening faults,
seeme to be relatively poor in volcanoes as compare
with areas such as the Pacific Ocean, where pressure
was increasing. The fronts of moving blocks mae
conitions more favorable to volcanism than i the
backs. Nevertheless, Wegener wonere whether the
mi-Atlantic rige might be consiere as a zone
where, with the continuing expansion of the Atlantic,
the floor was continuously breaking up, making room
for fresh, relatively flui an high-temperature Sima
from below! Moreover, increase volcanic activity in
some perios of Earth history might be ue to large
isplacements (as, for instance, uring the Tertiary).

Trench faults (Grabenbru che), i.e., rift valleys,

acquired new meaning as representing the beginnings
of new continental separations. Gravity measurements had shown that beneath such lines lay material
of greater density, compared to that on either side.
Therefore, these lines could be seen as incipient fissures within the continental blocks (into which the
denser Sima was rising according to the principle of
isostasy). The best examples of such separations were
provided by the East African trenches and their continuation through the Red Sea. At the majority of the
trenches, the measurable mass deficit was not compensated by greater density of the matter beneath it.
Thus, the trenches must be youthful disruptions of a
continental block.
Wegeners theory of mountain formation was further supporte by the fact that the foling of the
Anes seems to have been essentially simultaneous
to the opening of the Atlantic Ocean. The American
blocks, uring their westwar rifting, ha encountere resistance at the presumably very ol an relatively rigi floor of the Pacific Ocean. Thus, the
extene shelf, with its mighty seiments, forming
the western borer of the continental block, was
compresse to a range of fol mountains. For the
Tertiary fols of the Himalayas, Wegener assume
that lower Inia ha forme an extene peninsula
prior to compression, the southern en of which lay
next to that of South Africa. The fols ha been
prouce by impact of the Inian subcontinent an
the main mass of Asia.
Geological an Palaeontological
Evience
The palaeontological evience inicating a former
connection between the organic components of ifferent continents ha alreay given rise to the octrine
of former lan briges. Among the most striking
finings were the istributions of the Glossopteris
flora on the southern continents an the occurrence
of Mesosaurus at the turn of the Permian an the
Carboniferous exclusively in south-eastern South
America an the western parts of Africa; both of
these iscoveries suggeste a former connection of
the two continents. Using these relationships also
allowe calculations of when the continents were
separate (either by horizontal isplacements or by
sinking of the lan briges). South America an Africa
ha been connecte uring the Mesozoic, but were
separate at the en of the Eocene or Early Oligocene.
The connection between Europe an North America
seeme to have been maintaine uring the oler
Tertiary perio, but separation occurre in the
Miocene, although it might have continue in the far
north (over Scaninavia an Greenlan) into the
Pleistocene. The connection of Lower Inia with
southern Africa, which Wegener ha postulate
base on his ieas on the formation of the Himalayan
range, was also confirme by palaeontological evience. Zoogeographers ha long assume a former
elongate InianMaagascan peninsula (calle

Lemuria), separate from the African block by the

Mozambique Channel.
The zoogeographic concept of Lemuria ha given
rise to Suess notion of a great southern continent, Gonwana, comprising parts of South America,
Africa, Lower Inia, Australia, an Antarctica.
Assuming the unchange positions of its present-ay250 FAMOUS GEOLOGISTS/Wegener
relics, however, require ascribing a huge extent to
this continent. Wegener, by contrast, propose a
much reuce primeval continent, Pangaea. In the
Permian, i.e., until some 300 Ma ago, all the continents were supposely joine in one lan mass
extening from pole to pole. During the Triassic,
about 200 Ma ago, Pangaea began to break up an
the newly emerging continents starte moving into
their current positions. In the Jurassic, there were
few remaining connections except at the northern
an southern ens. Just as northern Europe an
North America remaine connecte until the oler
Tertiary perio, a connection of the southern continents seems to have persiste, running from the
southern coast of Australia over Antarctica to South
America. Later, the Antarctic block, like the
South American block in the Tertiary, move over
from South Africa towars the sie of the Pacific
Ocean. Only in the Quaternary perio, then, i the
Australian block become etache (Figure 4).
For geological an tectonic evience, Wegener
referre particularly to Suess magnum opus, publishe in three volumes uring 18851909, Das
Antlitz er Ere (The Face of the Earth). Consiering
the tectonic relations, Europe/Africa an both Americas seeme to represent the eges of an immense
expane fissure. In the north, for instance, the Greenlan massif was matche by Scaninavia, both consisting of gneiss, an the less mountainous North
America correspone to the likewise less mountainous Europe. The most striking example, however, was
the Carboniferous mountain range, calle the Armorican mountains (Suess transatlantic Altaies),
which mae the coalfiels of North America appear
to be the irect continuation of the European ones.
Wegeners theory of mountain formation was also
confirme by remarkable ifferences between the
Atlantic an the Pacific hemispheres, such as the
istinction between Pacific an Atlantic types of
coasts (marginal chains an ocean trenches in front
Figure 4 Wegeners reconstruction of the separation of the continents from the pri
meval Pangaea, from his 1926 paper Pala ogeo
graphische Darstellung der Theorie der Kontinentalverschiebungen , showing the r
elative positions of the continents during the Upper
Carboniferous (Jung Karbon), Eocene (Eozan), and Lower Quaternary (Alt Quartar)
(in two different projections). Cross hatching
represents deep seas, dotted regions represent shallow seas; rivers, recent coas
tlines, and outlines are shown only for orientation.FAMOUS GEOLOGISTS/Wegener 25
1
of the Pacific coasts, as contrasted to the wild, irregular ria Atlantic coastlines). There were also ifferences in the volcanic lavas of the two hemispheres, as
emphasize by the Vienna petrographer Frierich

Becke (18551931) an others. The Atlantic lavas

containe a greater proportion of soium, whereas
calcium an magnesium prevaile in the Pacific
lavas. Such ifferences were intelligible accoring to
the assumptions of continental movements. The
opening of the Atlantic was matche by the general
pressing of the continents against the region of the
Pacific Ocean: pressure an compression prevaile
at the coasts of the latter whereas tension an
splitting occurre at the latter.
Palaeoclimatology
Traces of glaciation uring the Permian (groun moraines lying on scratche berock) were to be foun
on the southern continents, e.g., in East Inia an
Australia. If the present-ay arrangement of the lan
masses ha prevaile at that time, this Permian ice
age woul have require an icecap of seemingly impossible size. An the north pole woul have been in
Mexico, where no trace of glaciation uring that
perio was recore. Following the iea of horizontal
isplacements, however, all regions subjecte to
glaciation came together concentric to the southern
margin of Africa. An one ha only to place the south
pole in this much reuce glaciate area to give the
Permian ice age a much more plausible form.
Wegener ha iscusse these palaeoclimatological
features since 1912. In 1924, he gave a etaile
escription of the climatological changes from the
Carboniferous through to recent times, following the
traces of glaciations, swamps, an eserts, i.e., moraines, coal, salt, an gypsum, throughout Earths
history (Figure 5). In reconstructing the respective
polar shifts, Wegener emphasize that they obviously
took place along with the great isplacements of the
continental blocks. In particular, there was temporal
coincience of the best confirme polar shift, in the
Tertiary, an the opening of the Atlantic (Figure 6).
Movement of the poles since the Pleistocene might also
be relate to the final separations of the continents in
the north an the south.
Motive Forces
Wegener was very cautious about the forces that might
have cause continental isplacements. First, it was
necessary to emonstrate the reality an the manner of
the isplacements before inulging in the hope of fining their cause. Nevertheless, he tentatively suggeste
two caniates: centrifugal forces cause by the rotation of Earth an tial-type waves within Earth, generate by the gravitational pull of the sun an the
moon. In the 1929 revision of Wegeners theory in
Figure 5 Wegener thought continental rift was the key to the climatic changes 
uring Earths history. This map, publishe in the
1924 book by Koppen an Wegener, Die Klimate er geologischen Vorzeit , shows tr
aces of glaciation, swamps, an eserts for the
Carboniferous. E, Traces of glaciation; K, coal; S, salt; G, gypsum; W, esert s
anstone. Dotte regions inicate ari areas, ashe
lines inicate the positions (i.e., the pathways) of the poles, an the bol cur
ve line inicates the respective position of the equator.252 FAMOUS GEOLOGISTS/
Wegener
Figure 6 Map publishe in the 1924 book by Koppen an Wegener, Die Klimate er g

eologischen Vorzeit , showing polar shifts (ashe

lines) from the Carboniferous to recent, relate to the African table (left, sou
th pole; right, north pole). Bol lines outline the continental
blocks; hatche lines represent the Carboniferous (Karbon) perio. Perm, Permian
; Jura, Jurassic; Trias, Triassic; Kreie, Cret
aceous; Eozan, Eocene; Miozan, Miocene; Beginn es Quartar, beginning of the Qua
ternary.
Die Klimate, he also mentione convection currents
within the Sima; these ha been first iscusse as a
cause of mountain formation by the Vienna geologist
Otto Ampferer (18751947) in 1906.
Wegener also eneavoure to calculate the recent
velocity of the relative motion of the continents, though he was well aware that these values must be
quite uncertain. In his 1912 paper, comparing various
longitue eterminations for Greenlan, he ha
euce an increase of the istance to Europe of
11 m year 1 . Referring to the lengths of transatlantic
cables, he suggeste that North America was rifting
away from Europe at about 4 m year 1 .
From Continental Drift to
Plate Tectonics
The theory of continental rift was long rejecte by
the majority of geologists. Among Wegeners few
followers were the South African Alexaner Du Toit
(18781948), for whom continental rift provie
the best explanation of the close similarities between
the strata an fossils of Africa an South America,
an the Swiss geologist E mile Argan (18791940),
who saw continental collisions as the only means of
proucing the fole an buckle strata he ha observe in the Alps (see Famous Geologists: Du Toit).
Nevertheless, Wegeners explanation of the PermoCarboniferous ice age impresse even his critics.
Wegeners reputation as a meteorologist an a
polar explorer contribute to keeping his theory
alive. His work was immeiately remembere when,
aroun 1960, surprising ata were obtaine from the
ocean floor: palaeomagnetic patterns alongsie the
mi-ocean riges clearly suggeste the spreaing of
the seafloor. Within about two ecaes, Wegeners
principle of horizontal isplacements of parts of
Earths crust became almost universally accepte, although, ironically, the process still lacke a consensus
as to its causes, though convection currents in the
internal mantle are most commonly avocate.
It shoul be note that Wegeners original concept
iffere from moern plate tectonics in essential
points, particularly with regar to the Sial an the
Sima. Accoring to moern theory, the (Sialic) continents o not plough through the (oceanic) Sima.
Instea, both continents an ocean floor are regare
as forming soli plates, floating on the asthenosphere, which, ue to tremenous heat an pressure,
behaves like an extremely viscous liqui (as Wegener
ha thought the Sima i). Therefore, the oler
term continental rift, still often use toay, is not
quite appropriate for the moern concept. Notwithstaning these ifferences, Wegeners basic ieas
remain soun, an the lines of evience that he

use to support his theory are still vali. He first

envisage a ynamic Earth, connecting its major
features an various geological processes continentalFLUID INCLUSIONS 253
movements, fole mountain ranges, rift systems,
earthquakes, volcanism, ocean transgressions, palaeoclimatological changes, etc. on a global scale. In
this sense, Wegeners theory was a true forerunner of
plate tectonics.
See Also
Africa: Rift Valley. Famous Geologists: Du Toit; Suess.
Gonwanalan an Gonwana. History of Geology
From 1900 To 1962. History of Geology Since 1962.
Palaeoclimates. Pangaea. Plate Tectonics. Tectonics:
Mi-Ocean Riges; Mountain Builing an Orogeny.
Further Reaing
Carozzi AV (1985) The reaction of continental Europe
to Wegeners theory of continental rift. Earth Sciences
History 4: 122 137.
Fritscher B (2002) Alfre Wegeners The origin of contin
ents, 1912. Episoes 25: 100 106.
Jacoby WR (2001) Translation of Die Entstehung er Kon
tinente, Dr Alfre Wegener, Petermanns Geographische
Mitteilungen, 58 (1912). Journal of Geoynamics 32:
29 63.
Ko ppen V and Wegener A (1924) Die Klimate der geolo
gischen Vorzeit. Berlin: Borntra ger.
Lu decke C (1994) Stratigraphische Methode der Rekon
struktion von Expeditionsergebnissen am Beispiel des
Todes von Alfred Wegener wa hrend der Gro nlandexpedi
tion (1930 31). In: Fritscher B and Brey G (eds.) Cosmo
graphica et Geographica: Festschrift fu r Heribert M.
Nobis zum 70. Geburtstag, Algorismus, vol. 13,
pp. 347 367. Munich: Institut fu r Geschichte der
Naturwissenschaften.
Oreskes N (1999) The Rejection of Continental Drift:
Theory and Method in American Earth Science. New
York and Oxford: Oxford University Press.
Runcorn SK (ed.) (1966) Continental Drift. New York and
London: Academic Press.
Schwarzbach M (1986) Alfred Wegener: The Father
of Continental Drift. Madison. WI: Science Tech
Publications.
Sengo r AMC (1991) Timing of orogenic events: a persistent
geological controversy. In: Mu ller DW, McKenzie JA,
and Weissert H (eds.) Controversies in Modern Geology:
Evolution of Geological Theories in Sedimentology,
Earth History and Tectonics, pp. 403 473. London:
Academic Press.
Wegener A (1912) Die Entstehung der Kontinente.
Petermanns Mitteilungen aus Justus Perthes Geogra
phischer Anstalt 58: 185 195, 253 256, 305 309.
Wegener A (1926) Pala ogeographische Darstellung der
Theorie der Kontinentalverschiebungen. In: Dacque E
(e.) Pala ogeographie, pp. 171 189. Leipzig and Wien:
F Deuticke.
Wegener A (1971) The Origin of Continents and Oceans.
(Translation from the 4th revised German edition by
J Biram, with an introduction by BC King.) London:
Methuen.
Wegener A (1980) Die Entstehung der Kontinente und

Ozeane. (Reprint of the 1st and 4th editions, edited by

A Vogel.) Braunschweig: Vieweg.
Wegener E (1960) Alfred Wegener: Tagebu cher, Briefe,
Erinnerungen. Wiesbaden: Brockhaus.
Wutzke U (1998) Kommentiertes Verzeichnis der schriftli
chen Dokumente seines Lebens und Wirkens, Berichte
zur Polarforschung 288. Bremerhaven: Alfred Wegener
Institut fu r Polar und Meeresforschung.
FLUID INCLUSIONS
A H Rankin, Kingston University,
Kingston-upon-Thames, UK
2005, Elsevier Ltd. All Rights Reserved.
Introduction
Fluid inclusions are small droplets of fluid that have
been trapped within crystals either during primary
growth from solution or at some later stage, usually
as a result of recrystallization along healed microfractures. They are ubiquitous in both naturally
occurring minerals and in laboratory-grown crystals.
To the chemist or materials scientist, these gross
defects cause endless obstacles in their quest to grow
near-perfect crystals. However, to the geologist, they
provide a unique fossil record of the various
fluids responsible for the formation and evolution of
rocks and minerals throughout the history of the
Earth.
Despite their small size (usually less than 20 mm),
their chemical composition and physical properties
can be readily determined, and the data may be used
to estimate the temperatures, pressures, and physicochemical nature of the fluid at the time of trapping.
This information has made an immense contribution
to the development of modern theories of ore genesis,
petrogenesis, diagenesis, and petroleum migration
and accumulation, and to our understanding of the
importance of the fluid phase in a wide range of
geological processes.254 FLUID INCLUSIONS
Occurrence and General
Characteristics
Formation and Genetic Classification of Fluid
Inclusions
Small changes in the chemical or physical properties
of fluids near to a growing crystal face can lead to
perturbations in the stability of crystal growth and the
development of gross defects, manifested as embayments, along crystal faces. These embayments will
seal over during a period of greater stability, trapping
a portion of fluids to form primary (P) flui inclusions. In many instances, the trappe flui will be
homogeneous at the time of trapping. In others,
where immiscible fluis are present or where mechanical entrapment of other coexisting crystalline phases
has occurre, trapping will be heterogeneous.
At some stage after primary growth, seconary (S)
flui inclusions can form from later fluis, particularly
as a result of recrystallization along microfractures.
The chemical an physical properties of these inclusions may be very ifferent from those of the earlier
mineral-forming fluis. However, if fracturing an
rehealing take place uring primary growth, the fluis

may be inistinguishable, an the terms pseuoseconary or primaryseconary (PS) appropriately

escribe such inclusions. A schematic representation
of this genetic classification of inclusions is shown in
Figure 1.
For most geological applications, it is necessary to
establish whether the inclusions are primary, seconary, or pseuoseconary, an also whether heterogeneous trapping has occurre. Heterogeneous trapping
may be recognize by the variable proportions of
liquis an solis in a single group or generation of
inclusions. Various criteria may be use to istinguish
between P, PS, an S inclusions, but these may be
ifficult to apply an it may be ifficult to ientify
primary inclusions in many samples.
Figure 1 Schematic representation of the istribution of pri
mary (P), seconary (S), an pseuoseconary (PS) flui inclu
sions in a quartz crystal. Moifie from Rankin AH (1989) Flui
inclusions. Geology Toay 5: 21 24.
in turbi or translucent minerals, such as felspar.
Quartz is usually the preferre host.
Size an Shape of Inclusions
Choice of Material for Stuy
The successful application of flui inclusion stuies
epens partly on serenipity an partly on the type
an quality of material available for stuy. Due to
their small size, observations on flui inclusions are
carrie out uner a microscope using polishe wafers
aroun 12 mm thick. In most cases, clear, transparent minerals are neee, but it is also possible to stuy
inclusions in some eeply coloure, semi-transparent
minerals in very thin (<50 mm) polishe sections.
Care must be taken with soft, easily cleave minerals,
such as calcite an fluorite, because of the possibility
of leakage uring sample preparation or analysis.
Flui inclusions are particularly ifficult to observe
Flui inclusions selom excee 1 mm in size; most are
less than 20 mm, an those greater than 1 cm are
exceptionally rare an usually regare as museum
specimens. Flui inclusions isplay a variety of
shapes. They may be flattene an irregular, roune,
or regular with three-imensional negative crystal
shapes mimicking the crystal symmetry of the host
crystal (Figure 2).
Phases Present at Room Temperature
Flui inclusions contain varying proportions of liqui
(L), soli (S) an gas (G) epening on the composition (X), temperature (T), pressure (P), an volume
(V) of the enclose flui at the time of entrapment.
humanity," he wrote, "one way that we can fin a purpose is to stuy the univers
e by the
methos of science, without consoling ourselves with fairy tal
Niels Bohr, a Danish physicist an leaer of this revolution, once sai that a p
erson who
was not shocke by quantum theory i not unerstan it.
This week, some 700 physicists an historians are gathering in Berlin, where Pla
nck
starte it all 100 years ago, to celebrate a theory whose meaning they still o
not
unerstan but that is the founation of moern science. Quantum effects are now

7invoke to explain everything from the perioic table of the elements to the ex
istence of
the universe itself.
Fortunes have been mae on quantum weirness, as it is sometimes calle. Transis
tors
an computer chips an lasers run on it. So o CAT scans an PET scans an M.R.I
.
machines. Some computer scientists call it the future of computing, while some p
hysicists
say that computing is the future of quantum theory.
"If everything we unerstan about the atom stoppe working," sai Leon Leerman
,
former irector of the Fermi National Accelerator Laboratory, "the G.N.P. woul
go to
zero."
The revolution ha an inauspicious start. Planck first regare the quantum as a
bookkeeping evice with no physical meaning. In 1905, Albert Einstein, then a pa
tent
clerk in Switzerlan, took it more seriously. He pointe out that light itself b
ehave in
some respects as if it were compose of little energy bunles he calle lichtqua
nten. (A
few months later Einstein invente relativity.)
He spent the next ecae wonering how to reconcile these quanta with the trait
ional
electromagnetic wave theory of light. "On quantum theory I use up more brain gre
ase
than on relativity," he tol a frien.
The next great quantum step was taken by Bohr. In 1913, he set forth a moel of
the atom
as a miniature solar system in which the electrons were limite to specific orbi
ts aroun
the nucleus. The moel explaine why atoms i not just collapse -- the lowest o
rbit was
still some slight istance from the nucleus. It also explaine why ifferent ele
ments
emitte light at characteristic wavelengths -- the orbits were like rungs on a l
aer an
those wavelengths correspone to the energy release or absorbe by an electron
when
it jumpe between rungs.
But it i not explain why only some orbits were permitte, or where the electro
n was
when it jumpe between orbits. Einstein praise Bohr s theory as "musicality in
the
sphere of thought," but tol him later, "If all this is true, then it means the
en of
physics."
While Bohr s theory worke for hyrogen, the simplest atom, it bogge own when
theorists trie to calculate the spectrum of bigger atoms. "The whole system of
concepts
of physics must be reconstructe from the groun up," Max Born, a physicist at
Gottingen University, wrote in 1923. He terme the as-yet-unborn new physics "qu
antum
mechanics."
Boy s Mechanics
The new physics was born in a paroxysm of ebate an iscovery from 1925 to 1928
that
has been calle the secon scientific revolution. Wolfgang Pauli, one of its rin
gleaers,

calle it "boy s mechanics," because many of the physicists, incluing himself,

then 25,
8Werner Heisenberg, 24, Paul Dirac, 23, Enrico Fermi, 23, an Pascual Joran, 23
, were
so young when it began.
Bohr, who turne 40 in 1925, was their father-confessor an philosopher king. Hi
s new
institute for theoretical physics in Copenhagen became the center of European sc
ience.
The ecisive moment came in the fall of 1925 when Heisenberg, who ha just retur
ne to
Gottingen University after a year in Copenhagen, suggeste that physicists stop
trying to
visualize the insie of the atom an instea base physics exclusively on what ca
n be seen
an measure. In his "matrix mechanics," various properties of subatomic particl
es coul
be compute -- but, isturbingly, the answers epene on the orer of the calcu
lations.
In fact, accoring to the uncertainty principle, which Heisenberg enunciate two
years
later, it was impossible to know both the position an velocity of a particle at
once. The
act of measuring one necessarily isturbe the other.
Physicists uncomfortable with Heisenberg s abstract mathematics took up with a
frienlier version of quantum mechanics base on the familiar mathematics of wav
es. In
1923, the Frenchman Louis e Broglie ha aske in his octoral thesis, if light
coul be a
particle, then why couln t particles be waves?
BALTIMORE, April 5 -- A gasp went through the auitorium of the Space Telescope
Science Institute on Wenesay when Dr. Aam Riess, a young astronomer from the
institute, put the last mark on his so-calle Hubble iagram, a plot of the brig
htness an
spee of istant objects that astronomers use to ivine the history of the unive
rse.
The Darth Vaers of astronomy ha gathere here to take stock of their expaning
an
increasingly ark realm. Once upon a time astronomy was about what coul be seen
in
the sky, about jewel-like lights that move in eternally recurring patterns an
the soft
glow of galaxies an comets.
Now it is about what cannot be seen. In the last few ecaes astronomers have ha
 to
confront the possibility that stars an galaxies -- not to mention the creatures
that inhabit
them -- are barely more than flecks of froth on a stormy sea of ark matter.
Now Dr. Riess was presenting his colleagues with evience, base on observations
of a
star that exploe 11 billion years ago, that the universe -- ark matter an al
l -- was
being blown apart uner the influence of a mysterious antigravitational force kn
own only
as "ark energy."
"We are oing astronomy of the invisible," amitte Dr. Mario Livio, a theorist
at the
Space Telescope institute, who ha organize the meeting, calle "The Dark Unive
rse:

Matter, Energy, an Gravity" last fall.

As it turne out, the meeting coincie with a NASA news conference announcing t
he
breakthrough iscovery by Dr. Riess an his colleagues an thus was ominate by
iscussions of new telescopes in space an new imensions in the universe as
astronomers grapple with the meaning of ark energy an how to take its measure
.
Now physicists, some of whom have been reluctant to take acceleration of the uni
verse
seriously, will have to explain what this ark energy is. "Those numbers are ala
rming,
an apparently true," sai Dr. Michael Dine, a theoretical physicist from the Un
iversity of
California at Santa Cruz. He escribe his colleagues as now working "franticall
y" to
fin an explanation.
19On one level, the universe, with all of its ark baggage, seems to make sense.
The total
amount of matter an energy seems to be just enough to guarantee that the largescale
geometry of space-time is "flat," or Eucliean, a result that cosmologists have
long
consiere to be the most esirable an aesthetic. On the other han, the etail
e
breakown of the constituents of the cosmos is, as Dr. Livio says, "ugly" -- 65
percent
ark matter, 30 percent ark matter of unknown nature an only 5 percent stars,
gas an
ust.
"We live in a preposterous universe," sai Dr. Michael Turner, an astrophysicist
at the
University of Chicago. "Dark energy. Who orere that?"
Of course, it was Einstein who originally orere ark energy when he inserte a
fuge
factor calle the cosmological constant into his gravitational equations escrib
ing the
universe. Lamba, as it is known, after the Greek letter, represente a sort of
cosmic
repulsion associate with space itself that kept the cosmos from collapsing of i
ts own
weight. Einstein abanone the cosmological constant when it was iscovere that
the
universe was expaning, an resiste efforts to bring it back, once referring to
it as his
biggest bluner.
But he couln t keep it out forever. In 1998 two competing teams of astronomers
trying to
measure how the expansion of the universe was slowing own because of cosmic gra
vity,
foun that the universe was actually speeing up, as if the galaxies were being
pushe
apart by a force -- ubbe, in the spirit of the times, "ark energy."
"This was a very strange result," recalle Dr. Riess, who was a member of one of
the
teams. "It was the opposite of what we thought we were oing." Was this Einstein
s ol
cosmological constant, something even weirer or a mistake?
The effect ha showe up as an unexpecte imness on the part of certain exploi
ng stars
known as supernovae that the astronomers were using as so-calle stanar canle

s,
objects whose istance coul be gauge from their apparent brightness. The astro
nomers
euce that these stars were farther away than they shoul have been in an even
ly
expaning universe, an that therefore the expansion was actually accelerating.
But ust or chemical changes over the eons in the stars coul also have imme t
he
supernovae. The cleanest test of ark energy an the acceleration hypothesis, Dr
. Riess
explaine, woul be to fin supernovae even farther out an back into the past,
halfway
or more back to the Big Bang itself. Because it is space itself that provies th
e repulsive
push, accoring to Einstein s equations, that push shoul start out small when t
he
universe is small an grow as the universe expans. Cosmic acceleration woul on
ly kick
in when lamba s push got big enough to ominate the gravity of orinary matter
an
energy in the universe, about five or six billion years ago. Before then the uni
verse woul
have been slowing own, like a Mark McGwire blast that has not yet reache the t
op of
its trajectory, an a supernova glimpse at that great istance woul appear rel
atively
brighter than it shoul. If ust or chemical evolution were responsible, such i
stant stars
shoul appear relatively even immer.
20By chance the Hubble Space Telescope ha observe a supernova in late 1997 an
early
1998 that prove to be 11 billion light-years away -- the most istant yet seen.
On Dr.
Riess s Hubble iagram it appeare twice as bright as it shoul.
"Extraorinary claims require extraorinary evience -- I hope the I.R.S. oesn
t say that
to you," Dr. Riess tol his auience, but, he conclue, "the cosmological const
ant looks
goo for this supernova."
Dr. Livio sai, "A year ago probably a large fraction of the people in this room
woul not
have believe it."
But there were more complicate explanations, forms of ark energy other than th
e
cosmological constant on physicists rawing boars, as well as the possibility
that
astronomers were still being foole. To explicate the nature of the ark energy,
astronomers nee to observe more supernovae as far back as 11 billion years ago,
to
cover the time when the universe began to accelerate.
"How fast i it go from eceleration to acceleration?" aske Dr. Riess. Answeri
ng such
questions coul help astronomers etermine how har the ark energy is pushing o
n the
universe compare with the preictions for the cosmological constant. A fast tur
naroun,
he sai, "begins to tell you there is a lot of oomph for a given amount of it.
"
"The cosmological constant is the benchmark oomph," he sai.
To fin those supernovae so far out cosmologists will have to go to space, sai

Dr. Saul
Perlmutter, a physicist at the University of California s Lawrence Berkeley Nati
onal
Laboratory an a veteran ark energy hunter.
On the groun, the supernova researchers have to employ a wie network of people
an
telescopes to etect the explosions, iagnose their type an then to watch them
fae. Dr.
Perlmutter escribe an orbiting telescope that woul perform all three function
s. The
Supernova/Acceleration Probe, or SNAP, woul combine an 80-inch iameter mirror
(only about 16 percent smaller than the Hubble), a giant electronic camera with
a billion
pixels an a special spectroscope.
If all goes well, Dr. Perlmutter sai, the telescope coul be launche in 2008.
In three
years of operation, he estimate, SNAP coul harvest about 2,000 supernovae. To
istinguish the ifferent ieas about ark energy, observations woul have to be
refine
to the level of 1 or 2 percent uncertainty .
"We re all excite," he sai.
So are the physicists. Their list of suspects begins with Einstein s cosmologica
l constant,
but therein lies a problem. About the time that Einstein was abanoning it, quan
tum
mechanics -- the set of rules that govern the subatomic realm -- was establishin
g a
21theoretical founation for the cosmological constant. Accoring to quantum the
ory,
empty space shoul be foaming with temporary particles an their cumulative ener
gy
woul outweigh the matter in the universe, incluing ark matter, by 120 orers
of
magnitue -- that is, a factor of 10 followe by 119 zeros. At that level, the f
orce of the
vacuum woul either have crumple the universe or blown it apart before even an
atom
ha the chance to form.
The fact that the universe is in fact puttering along rather gently suggests tha
t there is
something funamental about physics an the universe that physicists still on t
know.
Dr. Steven Weinberg, the Nobel Prize-winning particle theorist at the University
of
Texas, has calle the cosmological constant "the bone in our throat." If the ar
k energy
really is Einstein s cosmological constant, then physicists have to answer quest
ions like
why it is so small -- roughly comparable, in fact, to the ensity of matter in o
ur own
epoch.
Lacking an answer so far, even from string theory, the mathematically aunting c
aniate
theory of everything, some theorists have resorte to a controversial an somewh
at
philosophical notion calle the anthropic principle, which, in effect, says that
physicists
nee to factor in their own existence when they think about the universe. Out of
all the
possible universes that can be imagine, this line of thinking goes, the only on

es in which
humans can fin themselves is one that is conucive to human life.
That means, Dr. Weinberg has pointe out, that the cosmological constant has to
be small
enough to allow time for galaxies an stars to conense from the primorial fog
before it
takes over an starts blowing the universe apart.
Dr. Alex Vilenkin of Tufts University in Massachusetts pointe out to the Dark U
niverse
auience that the universe was at its peak in making stars about five billion or
six billion
years ago, just about the time that ark energy an the matter ensity woul hav
e been
equal. Our own sun, some 4.5 billion years ol, was on the tale en of that wave
, an now
here we are. "Observers are where the galaxies are," sai Dr. Vilenkin. "A typic
al
observer will see a small cosmological constant."
Many physicists are uncomfortable with this line of reasoning, an they are seek
ing the
answer in ifferent class of theories known as quintessence, after the Greek wor
 for the
fifth element. Moern physics, note Dr. Paul Steinhart, a theorist at Princeto
n, is
replete with mysterious energy fiels that woul exhibit negative gravity. The t
rick, Dr.
Steinhart explaine, is fining a fiel that woul act like the ark energy wit
hout a lot of
fuging on the part of theorists.
"The observations are forcing us to o this," he sai. "Dark energy is an intere
sting
problem. Any solution is quite interesting."
One theory that capture the fancy of the astronomers in Baltimore was a moific
ation of
gravity recently propose by three string theorists at New York University: Dr.
Gia
Dvali, Dr. Gregory Gabaaze an Dr. Massimo Porrati. In string theory -- so nam
e
22because it escribes elementary particles as tiny vibrating strings -- the or
inary worl is
often envisione as a three-imensional islan (a membrane, or "brane" in string
jargon)
floating in a 10- or 11-imensional space. Orinary particles like electrons an
quarks
an forces like electromagnetism are confine to three imensions, to the brane,
but
gravity is not.
As a result, Dr. Dvali suggeste that gravity coul only travel so far through c
onventional
space before it leake off into the extra imensions, thereby weakening itself.
To an
observer in the traitional three imensions it looks as if the universe is acce
lerating. The
cosmological constant, in effect, he sai, is a kin of gravitational brane rai
n. "Gravity
fools itself," he sai. "It sees itself as a cosmological constant."
Dr. Dvali s theory was welcome by the astronomers as a sign that string theory
was
beginning to come own from its ivory tower of abstraction an make useful, test
able

preictions about the real worl. (In another string contribution, Dr. Steinhar
t introuce
a new theory of the early universe, in which the Big Bang is set off by a pair o
f branes
clashing together like cymbals.)
Afterwar Dr. Riess an Dr. Perlmutter presse Dr. Dvali on what they woul see
when
they looke out past the crossover point where gravity began falling out of the
worl;
woul the transition between a ecelerating universe an an accelerating one hap
pen
more abruptly than in the case of the cosmological constant? Dr. Dvali sai he h
an t
one any calculations, but he sai it was his "nave guess" that the crossover wou
l
happen more smoothly than in a lamba worl.
"I  love to see this guy o some Hubble iagrams," Dr. Riess sai.
Even if Dr. Dvali coul be coaxe into proviing a preiction, however, success
in
ientifying the ark energy was not guarantee to the astronomers. Calling himse
lf a
spokesman for the "cranky point of view," Dr. Steinhart pointe out that the of
tproclaime era of "precision cosmology" was boun to have its limits. Other
cosmological parameters, particularly the cosmic ensity of matter in the univer
se, were
not likely to be known well enough for even SNAP to untangle the moels in which
the
quintessence varie over time. Worrie that the overselling of SNAP coul sap
astronomers will to come up with new ieas, he sai, "We shoul try to make as
few
pronouncements as possible."
Dr. Turner refuse to be swaye from his "irrational exuberance." Appealing to t
he
astronomers prie, he urge them to be ambitious. "We have a chance to o funa
mental
physics here," he sai. "Let s see if we can crack this nut. Maybe we ll fall on
our faces.
Maybe cranky Paul is right.
"I still have a lot of youthful juices in my boy."
Copyright 2002 The New York Times Company
23Mysteries of the Universe
IMAGINARY TIME
Before the Big Bang, There Was . . . What?
By DENNIS OVERBYE
What was Go oing before he create the worl? The philosopher an writer (an
later
saint) Augustine pose the question in his "Confessions" in the fourth century,
an then
came up with a strikingly moern answer: before Go create the worl there was
no
time an thus no "before." To paraphrase Gertrue Stein, there was no "then" the
n.
Until recently no one coul atten a lecture on astronomy an ask the moern ver
sion of
Augustine s question -- what happene before the Big Bang? -- without receiving
the
same frustrating answer, courtesy of Albert Einstein s general theory of relativ
ity, which
escribes how matter an energy ben space an time.

If we imagine the universe shrinking backwar, like a film in reverse, the ensi
ty of
matter an energy rises towar infinity as we approach the moment of origin. Smo
ke
pours from the computer, an space an time themselves issolve into a quantum "
foam."
"Our rulers an our clocks break," explaine Dr. Anrei Line, a cosmologist at
Stanfor
University. "To ask what is before this moment is a self-contraiction."
But lately, embolene by progress in new theories that seek to unite Einstein s
lorly
realm with the unruly quantum rules that govern subatomic physics -- so-calle q
uantum
gravity -- Dr. Line an his colleagues have begun to ege their speculations cl
oser an
closer to the ultimate moment an, in some cases, beyon it.
Some theorists suggest that the Big Bang was not so much a birth as a transition
, a
"quantum leap" from some formless era of imaginary time, or from nothing at all.
Still
others are exploring moels in which cosmic history begins with a collision with
a
Inspire by e Broglie s ieas, the Austrian Erwin Schroinger, then at the Univ
ersity of
Zurich an, at 38, himself oler than the wunerkin, sequestere himself in the
Swiss
resort of Arosa over the 1925 Christmas holiays with a mysterious woman frien
an
came back with an equation that woul become the yin to Heisenberg s yang.
In Schroinger s equation, the electron was not a point or a table, but a mathem
atical
entity calle a wave function, which extene throughout space. Accoring to Bor
n, this
wave represente the probability of fining the electron at some particular plac
e. When it
was measure, the particle was usually in the most likely place, but not guarant
ee to be,
even though the wave function itself coul be calculate exactly.