AND

EVOLUTION
BY

G. R. DE BEER
•

•
 PREFACE
HE present essar is the Outeome of a simmering
T revolt which has occupied me ever since the dis-
illusionment which followed the realization th-:u the
theory of recapitula.tion was not all that it W':\S cbimed
to be. The faith reposed in this theory I believe to be
ill founded, and, although I had originally no inten-
tion of sh3.k.ing that of others when I committed
these arguments to paper for my own use, I have
been tempted to try. Of originality of material this
essay claims little, and its function is r.uher to :l.tternpt
to replace on the rails laid by von Baer the train of
biological thought which was shunted off them by
Haecke!. The time for this seems to be ripe, for, as
my bibliography will show, the last ten years have
witnessed the publication of a number of works bear-
ing on some aspect of the problem, interest in which
seems to be reawakening. Should my humble effort
only raise hostile comment it will have succeeded in
focusing attention on this fuscinating subject.
I am indebted to my friends, Mr. H. W. Garrod
for assistance in interpreting a passage in Harvey's
works, and to Professor E. S. Goodrich, F.R.S., .
Priote:d in Great Britain Mr. E. B. Ford, and Mr. R. Snow for reading
 )'REFAC£ ~
VI • To Professor \V. Garst:lIlg I am
h m:llI11 scn pf. d I . h
Ie· h I f 1 suggestions,:lIl WIS to
fut for e P II CONTENTS
grate. . I k vledgemenr of the help which
de SptCl:l ac nO\ . • ••
m R
Dr E. S. ussr:l.,
ll'~ book Form and
. Fun{/Ion, has List of l1lustrations . . . VIII

. • me m
glyen . fu CITtating
I the findmg of references I. Stages of Development and Stages of
and other m:me.rs. Evolution . 1
G. R. DE B.
I r. Ontogeny . 12
MuTOS COLLECE, II I. Speeds of the Processes of Development 19
OxrORD.
IV. Phylogeny. 28
Srpumhtr 19'29·
V. Heterochrony and its Effect in Phylo-
geny 34
VI. Caenogenesis • 4°
VII. Deviation 4S
VIII. Neoteny 57
IX. Vestigial Structures due to Reduction 71
X. Adult Variation
73
Xl. Vestigial Structures due to Retardation
75
XII. H ypermorphosis
76
XIII. Acceleration 80
XIV. Paedomorphosis and Gerontomor-
phosis
• 88
XV. Repetition
• 101
XVI. Conclusions
• •
XVII. Bibliography 1°7
Index 1°9
• • • • 114
 LiST OF I LLU TRATIONS
I
I The youns and adult (orlllS of three ~jc,. to aho w lIlt
. . 'l~rilr between the .dulls Ilnd the differences betwcc STAGES OF DEVELOPMENT AND
Ilml M' II 'T'L At n
STAGE OF EVOLUTION
the )'oung. Afler L. ',. lit , .l"~ H~/UrQI JIiJ/o'] oj
Afroli( /tUUIJ, by pcrllm"on of MacmIllan & Co., Lid. U

2. An embryo of a chick ~ol~lpa.rcd with that of a human

being to ahow the slInllllfllY between lhe young of
T HE life of an animal may be said to start from
the egg which has just been fertilized. The egg
is a more or less spherical object and it bears no
animaIJ whose adults arc markedly diffcrcni. After llal.
dane.nd lIuller. A"i"'QllJi%ty, and by ~rmiS$jon of resemblance to the animal that laid it, or into which
ProCOIOr Arthur Thomson 46, i7 it will develop. The processes of deve1o~ment are
therefore concerned with the transformatIon of the
}. SUgtSin thedevdorment ofthe limpet. From W. Gantang.
TItt Orililf tlftd EDU/.I;," of LIlfT,'1l1 Fl1rmJ, by permission egg into that form which we recognize as 'the
of the Britj~h A.ocinion for tile Advanctmcnt of anima, I' . The transformation is gradual and SOI' the I
Science • 53 shape of the spherical egg is modified little .b>' ttt e
4· Thungk which the head makes wilh the trunk in embryo into another shape, and this is changed mt~ yet
and aduh dog and hum.n being. From L. Bolk, Pro-- different ones which succeed one another u~ul t~e
Iltlllt ur Mtllumctrllwg, 1926 (G uSlav Fischer, Jena) 61 definitive or adult shape is reached. Du;g I(~
S· A comparison bet\\een the larva o( an Echinoderm and development therefore, the animal passes roug f
thtform of alypical primitive h rdale. A(ter W. Cn- ' . stages, an d it is the sequence
a series of successive k of the0
u.ng, in QlllrltrlJ J 'IIr"al If MicrtJlCopifgl Sciclru, vol. which we mean when we spea
72, by pc.rmiuion of the t.:ditors. . . 65 these stages f h togenetic stages are
6. The brral r. r' ' 'I' 'J Some 0 t eon
anima s on/ogen . c Ie as the cater-
orm 0 I 1\lynlpod lOOn a(ler hatching. From .. such lor examp ,
','" ,RUb, IJtr;t;/t Jrr Nttlllrr"Uhelrdo, Geul/uhujt %II familiar to everyone, , Ie of the frog. But on
•
"'1,,'0 I J'
'I ,Vo ..... 189t(PauISic:bed.) . . 68 pillar of the moth or the tad'7' t of anim:'lls are
7· I'becrownlofth the whole the stages of deve opmen and he must
Frorn W K e IIppcr ,molars of embryos o( Ihe Dugong. to the layman,
Ntlll~" ItIl Ohmhal In Drduhrifun aer MediciwiJfh- morc or less un k"nown b "ness it is to col·
II It;..tJlirh C ,. b logist whose USI • I'
(C Ulli ... F' l rll rJr/luhafl ZU leWD, vol. VtJ rely on the em ryo, '. hases of an anIma s
lkuer, lena) • , 86 lect and study these mterestlllg P
existence. can be made alit from
Another series of stages
.
 : STA GES OF DEVELOPMENT AND
" b paring the full-grown or adult shapes STAGES OF EVOLUTION
anJOlals Ycom . th f h k" d Serres: 'Man only becomes man after t--Ye " 3
k "
mOd f animal With ase 0 at er In 5, and " . .~ ~Ing
ofone d" d tranSItional orgam2atory states which assimilate him
" th' they can be arrange In an or er of in-
findmg a " S h first to fish, then to reptiles, then to birds and
"
creasmg or decreasing complexIty. ue an .order
mammals.' Similar ideas were also expressed b}.
was not un kn own to the Greeks, . who called • It the
Meckel.
1 ,I h(i,,~s. At the top of ArIstotle 5 scale Was
J(alt OJ bo' I" " According to these views, therefore, the ont~
man, below whom came wha es, . VIvIparous quad- genetic series and the scale of beings run parallel
rupeds (other m~~mals), o~parous quadrupeds
with one another, and the developmental stages of
(reptiles and amphibians), .and bIT.ds) and below these
an animal are held to correspond to the definitive
a medley of animals whICh Aristotle regarded as
or adult stages of animals lower down on the scale.
being without blood, and in which we can recognize ';Vhile the scale of beings was gradually being
what we call the invertebrates, consisting of octopus, amended and expanded as new animals came to the
lobsters, insects, snails, starfish, and sponges, in notice of naturalists, the foundations of an accurate
descending order. A couple of thousand years later knowledge of the ontogenetic stages through. whic~
the scale of beings received complete expression at animals pass in their development were not laId untll
the hands of Bonnet, who strung out all the animals, the work of von Baer. As a result of his extensive
plants, and minerals which he knew on a long ladder researches, von Baer came to a number of conclu-
of which they formed the rungs. sions which are expressed in the form of four state-
It w~s natural that the thought should arise of ments which have come to be known as the 'laws of
comparing the two series which we have now just von Baer' As they are of the greatest importance
0

seen-~he sequence of ontogenetic stages and the from the present point of view, they are here repeated:
successIVe shapes f " I " 1. In development from the egg the. general
Th IS Idea seems t0 amma
O 0

h
s on the scale of bell1gs.
characters appear before the speCial char-
h 0 ave occurred to Harvey for
w en speakin f d ' , acters. th I
steps whO h go evelopment, he says:' ature, by
IC are the . From the more general characters e ess
animal wh same 111 the formation of any '2. general and finally the special characters are
atsoever goe th
animals, as I mi ht s rough the forms of all
graduallYacq . g say egg, worm, embryo, and developed. . 1d rtsmore
Ulres perf; f . During its development an alllma ep~ I
same notion' ec Ion WIth each step.' The 3· and more frolll the form of other anIma s.
IS expressed' h r
111 t e lollowing passage of
 STAGES OF DEVELOPMENT AND
+
+. The young stages, in the development of an STAGES OF EVOLUTION
5
animal are not like the adult stages of other .a mere static row .of shapes of adult an'lm as, I an d,
animals lower down on the scale) but arc like lns~cad.) became a ltne of ancestors which by modifi-
the young stages of those animals. cation In descent have evolved into other (and usually
By his 1St and 2nd laws von Ba~r expr~ssed the more complex) forms. The modifications undergone
fact that in the development of the chIck, for Instance, by the shape of the adult animals in ~ucces~ive
generations during evolution could now be regardcd
there is a stage at which he could recognize it as a
as a series of stages in the history of the racc, which
vertebrate but could not say what kind of vertebrate
series became known as the phylogen}. The problem
it waSj and that at a later stage when he could recog_
now was to see how ontogeny and phylogeny wcre
niu it as a bird it was still impossible to distinguish related, and the next step in this direction was taken
which kind of bird, The 3rd and 4th laws express by Fritz Muller. For MUller, ontogeny could follow
von Bacr's most important contribution, which is one of two methods. During its development from
that animals are more similar at early stages of their the egg an animal might either pass through the
development from the egg than when they are full ontogenetic stages and beyond the final adult stage
grown, and that this resemblance between early of the ancestor ('overstepping'), or it might diverge
stages becomes progressively diminished as they more and more from the ontogenetic stages of the
grow older. Indeed, of a couple of embryos which ancestor (progressive deviation). The former mode
von Baer had preserved in spirit, he was unable to reRects the theory of parallelism of Harvef and
say ~h~ther they were reptiles, birds, or mammals, Serres the latter gives expression to von Baer's
so similar are the young stages of these animals. thear; of the greater resemblance between ~nimals
Instead therefo
J •
f .
re)o passmg through the adult stages when they are young. It is important to nO~lc~ that
of other animals d ' . Muller bases phylogeny on ontogen),) for It IS the
. I unng Its ontogeny, a developing
amma mOves a f , ontogeny ('I.e. aItera t'on 'In the processes
Baer d h way rom them, according to von changes In I .
Jan t eontog , of development of the descendants as compared with
to the s enetlc stages do not run parallel
equence of r. f those of the ancestor) which make the adult descen-
The introd . Orms 0 the scale of beings. . dsoaddanew
. UCtIOn of the f ' dants differ from their ancestors, an
obVlously dest' d concept 0 evolution was
Ine to m k . link to the phylogenetic chain. k I took
consideration of h a. e Important changes in the
fb' t erelatlon f This was the state of affairs when Haec e I '
o elngs. For th So ontogeny to the scale · . ws of the re atlon
e scale of b ' the matter up and expresse d h IS vie
etngs then ceased to be
 STAGES OF DEVELOPMENT AND
6 to ph),logeny in the form of his famous
of ontogeny . ., . therefore, had no ancestral or historic
f ttapi/uln/lo" or !)loge/rellC law. AccordIng
theory 0 r. b d d h cance.
th' togeny is to e regar e as a SOrt re-
to . 1~, t~:n of phylogeny, and phylogeny is the
:~~~~~ical cause of ontogeny. The adult stages of
ancestors are repeated during the development
the
d e d back into
of t he descendants, but they are crow f;
earlier stages of ontogeny, there ore making the
latter an abbreviated repetition of phylogeny. These
repeated or 'recapitulated' stag~s reflect the history
of the race, and so Haeckel applied to them the term
palillgmttic. As an example may be mentioned the
stage in the unhatched bird and unborn mammal
when gill~slits or pouches are present. I-Iaeckel urged
that these gill-slits represented the gill-slits of the
adult stage of the ancestral fish, which in the birds
and mammals has been pressed back into early stages
of development.
At the same time, it could not be denied that in
so~e cases at least the young stages of a developing
antmal prese nt ed s hapes and structures which no
adult ancestor e Id 'bl
. ou POSSI y have possessed such as
the birth-me b f )
these cases H~ec~::s 0 la dmadmmal for inst~nce.. In
of p I' , COne u e that the recapItulatIOn
a mgenetlc sta
and that a new or ~;s was t~mporarily in abe~ance
calated' h t"ogenetIC stage had been lI1ter-
111 t e ontog . that the characters w IC appeare
•
rOung animal .
e;h'
Inevitable COnditions a~ an adaptation to some
Ie the mode of life of the
Imposed. The caenogenetic stagesJ
STAGES OF EVOLUTIO:N
7
signi6-
It will ~e no~ice~ that the biogenetic law abandons
von Bacr S princIple of progressive deviation , or
rather. relegates it to .rh: state of the caenogenc:tic
exceptions, and that It 15 really a reversion to the
f
II I'
theory 0 para e 15m and the 'overstepping' of
Muller, to which it adds the causational idea that the
succession of palingenetic stages in ontogeny is due
to these stages having succeeded one another in
phylogeny. The biogenetic law makes it necessary
to believe that the new variations (by means of which
evolution was brought about) occurred at the end
of the ontogeny of the ancestor, or in other W"ords
that the evolutionary novelty first appears in the
adult. Phylogeny, then, according to Haeckd, is
brought about by the successive tacking o~ new
final stages on to the existing adult stages of animals,
and the processes of development in ontogeny are
'due' to this progressive accumulation in phylogenr·
"'hat
vv 'd"e' may mean we shall see in the next chapter.
Seeming confirmation of Haeckel's views was pro--
vided by the work of Hyatt and "rurtemberger .on
Ammonites. These fossil animals had shells which
I dd d d ' g their life, The
were constant Y a e to unn
above-named authors thought that they could show
h' h d on the older parts
be
of the shell of an ancestral Ammonite were to
found on the younger parts of the shell of subsequent
_
•
d:tnt. nUllonit. t aJi events) this vie STAGES OF EVOLUTION
",d "cen •. h W inherited'. These t\~o authors th . . 9
-..! itself to \\ eIsmann, w 0 sunlmed
muteD dC'U I • up phylogeny to ontogeny b . m us pro~e to rdute
. . ' as follow·s: The onragen - ariSes fro . d ) ~s of the lnherinnc:e
hl5 opInion _. m
_I~ . bY !l con ens.'ttJ.on of Its stages') which o f acqmre characters. Quit f
the ph '0 . , . fth
ht' tl:- 6~ :l.s. ~ retra non 0 . e phyletic
. .
we accept or reject the mherita
e aput rom whethC:f
f .
nee 0 :tCqulrro chu-
:lcters (and we shall have something t bo .
hrlocrnt'ci) acqulSJtJO n S of ~e m~ture animal • . ' 0 S3Y 3. unhts
III the next chapter) It must be: born- ·In . 'th at t he
«Per-
. ,
llld deeper into the germlllal hIStory of the . ... nuou
rel:ltlon between ontogeny and phylo,-,cny is to be
spe<Jcs . • looked for and found in heredity. .
.~t the present day the theory of recapItulation . uch is the history of the de~'~lopment of thought
,:ill has its st2unth supporters. From the embryo- whIch has led to the theory of reo.pitulation which
logical point of \iew i\lacBride states that 'the larval still has :l remarkable sway. his respom;ible for the
(i.e. wly) phase of development represents a former fuct that every biological examination pJ.}--'(f ansv;erC'd
rondi ·on . the adults of the stock to which i, by :l schoolboy conmins the statement that 'during
bdon o '. Further, when comparing the 'oung stage its life-histor . an animal climbs up its f:unil)' trC'c'.
-nth 'the.dul, ancestral stage which it represents ... The criticism to which the biogenetic law 1nJ theory
on~ receives the impression that one is dealing with of recapitulation hayc been subjected seems to ha\-e
, reaction whith constand· repeated through thou· made but little impression, for the schoolbors'
sands, nlY myriads of generations) tends to sec in answers do not change, and yet th~ criticisms are
soo.ner an . SOOner in the course of deYeiopment; just not unimportant. In the first place it was pointed out
as Ul e life of the individual, the formation of habit (by Agassiz, Keibel, and 1ehnert 'mong othe...)
that the order in which chat.lcters appeared In ph .)()oo
Cluses reactions to require for their evocation less
and. less of th e ongtn3
.. I stimulus'. From another geny is not alwars faithfully reproduced in ontogeny.
pllln of \iew, Sm,·th \\'ood ward speakmg . For instance, teeth were e\·oh·ed before tongues, but
palaeonrolno;st sa' h h . I '.
as a in mammals now tongues de\-dop before te ~h. T
. h . -t::>o , )s t at e IS convmced that when- . an d revers..1 I a f the sequence
. alter-man ot Sf
e'ie:r e IS able to r thiS
.
.
I' d and It creates t
th, "ta· I. trace lOeages he finds evidence of the term htterochr01lJ IS app Ie , • h
pltU atlOn of a
hiSton-' d 'h' ncestra
I ch aracters m
. each I·'
uc- . . th II d etr'lCtion of adult c.
-J)3n elseq II . SuspICIOn that e a ege r . C·f· "'1"
m'n, h, L.. ua Ycon"nced that the phena- f d velopment I It rc<u I
o"",n·es h acters into younger snges 0 e .. f
<>plain,d b. w. en tracing lineages can only be . I d the pilll1g up 0 ne
occurs) is not sunp y ue to
l assumIng lh at ac U1eed. characters are jb66 C
 ' D£V£LOP~l ENT AND
S'rAG ES OF STAGES OF EVOLUTION
" h d of the life-history, for the order "
.' at t e ell ontogenies and so changed t~e shape of the final
\~Jrl:I!lOns.. t ecessarily respected. I t also sug-
stage of development ' viz
. ' the adult . Bu l aseneso
' f
or t hr rik .'IS nO n •
t legitimate to speak of a stage'
.
adult forms m~(hfied In this way is phylogeny, and
ts th'lt It \s nO .
ges 'of J b k to:l \:l.tcr or on to an ear her period so. phy.logeny IS the result of ontogeny instead of
beioU' sh, tC ;tC h" h' ,
, • \'r h' tO f )', It is not t c stage W leh 15 being Its cause. Garstang has thus arrived at the
' J 'I but certain characters W h'h
tn1hett"'IS
Ie may be same point of view as Hurst, who wrote: 'I do deny
Shlftc tlf l! Q{,
culiar 10 the stage. . that the phylogeny can so control the ontogeny as
pt Another line of attack was made possible by the to make the latter a record of the former.'
. se 'n detailed knowledge of the young stages. Among other recent critics may be mentioned
mere:!. I
Sedgwick showeJ that the earlier. stages of develop- Naef, who by his 'law of terminal alteration' redirects
ment of quite closely related animals (stich as the attention to von Baer's principle of greater resem-
hen and the d\ICk) could be distinguished, and blance between young stages; Sewernow, whose
W. His concluded that even at these early stages principle of 'anaboly' resembles the 'overstepping'
of MUlier, and whose 'archallaxis' savours of the
developing :mim:lls possess the characters of the
principle of deviation; Franz, who proposes four
class, order, species, :\l\d sex to which they belong,
methods ('biometabolic modes') by means of any of
as well ns individual characteristics. O. Hcrtwig
which ontogeny may be related to phylogeny; lastly
went on to point out that the very egg itself must have
Spath, whose conclusions are of particular interest,
specific characters although they may be invisible,
for they are based on a study of Ammonites, i. e. the
and thar the eggs of two different animals arc really same material as that which served for the develop-
as distinct from one another as are their adults, the ment of the views of Hyatt and \VUrtemberger. In
distinctions becoming more and more visible as this connexion Spath makes the following very im-
develo~ment proceeds. The egg of the mammal and portant statements: 'It may be necessary to assume
the u.mcellular ancestor arc not really comparable, an inverted geological order if our vie~vs of the
and In a .remark abl y t horough analYSIS " Hertwlg biological order of Ammonites are to contl.nue t.o ~
revealed thiS flaw'In the \oglc
' of Haeckel's argument. governed by discredited "laws" of recapItulation;
Recendy Garst
ski\full.' ang has elaborated this view very and 'when the horizons of all these slocks are
}, and has sh
evolutio \
h
Own t at there has been an definitely known it is hoped to get g~ :tddi-
n.onglhelin.eo ffertl'\'ned eggs (or z.ygotes)
in eonseq tional evidence for a final rejection of the views on
lienee of wh 1Ch ammals
. have modified thel. f
 I'H~I'\ nl' 111' \ 1",," 11"'1'1'
II '
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\\
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ncsiulI\ chl{)l'id(') :m: added to tht w,ltrr in wtmh :1
Shlulr it cll\lil'ol1l1\('l\t. N w the illll'rllal fflltors
\\rrr !'fCS(111'Il\ II 1(' fcrll'I'I/nl cp,g', :\IId ~O Ihey elll
riqh (I"undlll\l ) i developing, Iholt (, h willllrllh'r~()
o

n m difi d pro((' , of devclnpnltl11 and h,lvr 1\01 IWO

,I "heI :rrr'lrdrJ
"
I '
tiS t Ie.: tr.ln Illllied (,I tors, pa "llg'
" f \\hl~h fr III , tr
'yes but 11<\ ns Stotk.1rd f1hownL 'ountle, "nlilar
ill', Wh 1 arl'nt 10 ollspring l'tlll tilliteS IH.-red· examples Illig hi he given, hut Ihi. one ufhce w
llll('flli\\ f
tn, thefefor' ' l, W nil. k t I\t' qllc:~lt' II: I ) 0 I he show Ihal hy thl'm-;e1vr lhe intcrn.1 and lherefore
.lUOf whll.h 11 t 'U-·11l{ • a I\ormill
nrc pr 'Sellt in th· fl'rlili/ed ('A~ ,
j
sutlic c to ' t1'IlIlSllIIfICt! f,ldor nr nOI a' e U I ou
,
\HIIlI\,M \'
;jl,: QUill for II I f
\C 11 rl1\1l dcvd pili 'lit 0 :t11
, \r nrc nl~o q , , , . nniuHII. . I" ,llown Ih:llilI .IIHCr~
IS 1'(''\1)01 ~'II f UCSIIOIl1l1~ In how (:11" hcrc(ht)' NtlIOCI'OHf~ ('xp 'rJlIlcnf8 l,lve
_ __ 1.lll' Q I
' Se.UIII'III'" .~ 0 ••• , ...... n •• "'..... whi,c.I... _
 NT ,ENY
ONTOO I,NY
" '1 r. f nn :lllim:'\1 nl'C pos. eBsed equnlly hy the rO CBS of produ ti n in the ofT . .,
n:1 1Iott II P , pflnR (if (.h
crSII .I/l ,II
,
it! I,,"t.. S , lhe te ' 0 th . he,d 'of a similar to those whl(,;h wer po e, d I trt<trr,
w rill C"• I by rcgenenltlOn pr duce :l lall,. and the 'An organillm i. mould d It the rt \'ythflltrerll,
ell. of the "il can regenera,e a head. llutlf the cell. . h" \UtfJflhc
:lellOn between t e (;Ondillfmt tor 11m I' , Ihlrr~
of t he head possess the internal r.'CI rs. whi h ontrel
, ' U'Whl<h L
ul' 11S environment and lh f:Ulor f' ttla..e
Ihe produ tion of a tail a ~ell as Ihe I nlernal rauors
.
"
No slIlgle part
' I
I' h
,() Inhrru
completely acqui d
Tt', C1r duttl
.oIl\(t•
which entrel the pr ductl n of a head, how is it inheritance a one. araucrs are due t )
Ortp'JIlt
Ih,t in n rm,1 development the,e ell, do produ e I
and have to )e rna de anew at tvcry ,
" ' ~tntTiIotltjn '
a head ,nd not a tail? hild ha s Iv d thi, pr Ilem Similar views have been cx\)re d I r. 'I .
by ,howing experimentally lhat the first thing which ' I ' h' 'y '-Am .. In.
fhese con', u'uon" w Ich are htl cd On dcfi nl'I eexpc1'\_.
has 10 be sell led in a developing egg is the polarity, • mental eVidence, have a far-rtiuhinK Imp<Jfl1n(.c, In
i.e. which part of the egg will give ris 10 the front I he first pia e I hey Bhow thaI the qU,,"on-Are
and which t the hind end ofth future :lllim:'\1. N w acquired haratlcrs inh~ritc:d? hli' no mttning,
the determination of this polarity is th . result of the and lhat what the Cjuc6tloncrs really mean i.-(an
action of fact rs which arc external to I'he fertilized external faclors become internal? When, ,hrrt'fCire,
egg. Alilhe w'y thr ugh development the internal 1 fyatt defined an acquired eh, raCIer as', modifica·
faclor. produce no'hing f themselves, but 'hey tion which makes its appearantc in the adult or later
enable Ihe animal t rca t in definite ways to the stages of development and i. obviou Iy dep'ndenl
external factors and by this means give rise t struc- for its rigin upon other than hereditary taU " he
ture after stnlcturc in the pro css of development. i. really describing the e/Tect of an external factor,
Heredity docs not aCCOunt for the individual, but but that dOCB not justify him in excluding the parti.
merely for the potentialities some of which arc cipation f internal factors in producing that dtect,
r"llited in the individual. In ther words thc inter- We shall return below to the question as towhtthtr
na and transm' d f ' external fa tors may become internal, and may go on
' od' lUe actors arc by themselves unable
J tOpr uc' , now to note that the conc1usion5 at which Child and
• e an animal at all,
rhe same po
oodr' h
f'
'nt 0 VIew has been developed by oodrich have arrived lift a great burden off, lh~
bc dra~n' b
Who problem of heredity_ The structure of ,an anima,
strcsses the distinction which has to en •
h
SOWS a numl cr of exquisitely c lca
d I' to adJustm h
internal f ctwecn the process of transmission of Ihe
. actors fro
m parent fO offspring, and the
th e sp I·Inters inside a bone are situate
. d exactly w ere
 ONTOGE Y
~e' are required to withstand the pressure to which ONTOGENY
. " 17
th ) bone is subjected; the fibres of a tendon lie nerve-centre IR prOXimity to them l as Detwiler has
e d . along the line of strain between a muscle proved. The nerve-centres arise afresh in ea h
areur:ll ) .., h . d C
d h bone [0 which It IS attac ed; centres of generatlon l an by arising in a different place uod
111(e . del . er
nen~C'l
ls in the brain are .situate ase
.
to the ends the stimulation of a different set of nerye-fibres the
of the nerve-fibres from which they hablt~ally receive effect of a migration in phylogeny (or neurobio~xis)
.
unpu I~_
:>0.;>,
and when in phylogeny
"
there IS a change receives its explanation.
. the nerve-fibres from which any given nerve_ All that is required is the transmission by internal
:nrre habitually receives its impulses) the nerve_ factors of the capacity to react in these wa),s to new
centre migrates towards its ne".' ~ource of stimulation. stimuli which evoke one response after another.
It is obvious on looking back over the subject-
h is this change in the poslUon of nerve-eentres
matter of this chapter that ontogeny cannot be
during C'o-olution which Kappers has designated
regarded simply as an extrapolation into the future
under the term neurobiotaxis.
of a chain of events which happened in the past.
It was a source of much perplexity to the student Each ontogeny is a fresh creation to which the past
of heredity to try to understand how the factors contributes only the internal factors by the means of
controlling the production of such intricate and re- heredity. The action of the internal factors is to
fined adjustments could be transmitted to the fer- ensure that if the external factors are normal and
tilized egg and produce these effects. He need not do evoke any response in development and produce
h1\"e worried. about the production of the effects, for an animal at aliI that animal will develop along the
that lay in the province of the experimental embryo- same lines as its parent. The internal factors are only
~ and not in his. It is the very pressure in the a partial cause of ontogeny.
tiSSUes which causes the cells along the lines of stress \Ve may now turn to the questio~ as. to whether
10 od
~r uce sp I·Inters of bone, and that is why these the internal factors which are operative In ontogen.y
splinters are in the 'right' place; it is the very pull can in any way be regarded as the result ~f the mo:ch.
exerted On th·
.
b .
e tiSSUes y the muscles which deter-
fications which the ancestors underwent 10 evolution
tmnes the prod· f .. resulting in phylogeny; or) in other words, whether
. UctlOn 0 a tendon With Its fibres
onentated alo h I· phylogeny is in any way a 'cause' of ontogeny. If
ends f ng t e Ines of strain; it is the very
o the nerve-fib res W h·Ie h have the property a f these modifications which the ancestors underwent
stimul . were the result of changes in the internal factors,
atlng nerve-c II d··
e s to lVlde and so produce a
".. D
 UN j VVj",,~ •
,8
question can be answered at once, or rather
. d
h
thente .'
t arise for In thIS case the phylogeny will
It oes nO ) •
.I[se If have been the product .
of the Internal factors
working in ontogeny. But If, on the other hand, the
modifications which the ancestors. underwent in
pbylogeny were due to changes In the external
rs then these external factors could only become
fac t 0 ) .
the cause of the internal factors which control that
same modification in subsequent ontogenies in so
far as they can be converted from external into
internal factors (i. e. in old terminology, in so far
as acquired characters can become inherited I).
~ow the recent work of Heslop Harrison and of
Muller has shown that it is possible to produce a
permanent change in the internal factors as a result
of stimulating the animal with external factors) but
these external factors had not produced any modifica-
tion in development atall before they became internal.
Feeding moths on salts of manganese does not turn
their bodies hlack, although it can 'produce' an
mternal factor which will ensure that later onto-
genies will result in black animals. Harrison and
MU~ler have shown the possibility of 'direct induc-
[Ion ofacha . h .
f nge In t e Internal factors by external
I
actors. They h
ONTOGENY
·bl·
POSSI e, It seems t hat no case has )'et be 1j
·1 d . en satls-
facton y prove . In which the devdopme
nt 0 f an
ancestral ammal has undergone a modification as a
•
• result of external factors and in which th--..."'" ex t ernaI
factors have become internal and transmitted, so th
·fi . U
~ he same modI catton has been invariably produced
In all the subsequent ontogenies of dace-munt
animals in the absence of these external facton which
originally evoked tbe modification. ntil such a case
has been proved it cannot be ~lieved that the effects
of use and disuse on body or mind are transmittC'd,
and it cannot be stated that phylogenr pia).. an)· part
in ontogeny at all, even in the restricted part plaled
by the internal and transmitted factors.
Somatic induction, or the transmission of the
effects of use and disuse, constitute the kernel of
the Lamarckian point of view; and it is curious that
while we still want evidence of this, Lamarck ruled
out that of which we have evidence, viz. direct
induction.
III
THE SPEEDS OF THE PROCESSES OF
DEVELOPMEl\T
E have seen in the previous chapter that

facto
'.
somatic tnductio ,.
od
rs pr uced b
f
. ..
ave not shown the posslblhty of
n, I.e. 0 a change in the internal
W development was brought about by means of
. f·
the interactIon 0 mterna
I a d external factors.
n d
ain constant an
body whO h I Ya change in the structure of the Granted that the extern aI factors re m .
IC aUer ch . . . . to the way In
external fa t ange was Itself produced by normal, we have nOW to inquire 10
c ors. T o p-u t.h.e--lruttte.r~ C'le:arlv as
.' ,
":PEE.DS Of Dtn
the- inttn':oIf.t
~Lur 'I n;~ I

produce thoir meets,

, . __ I h:torS b2\-e recencl ' bttn subj=<d
SPEED OF DEYE.LOP. E.~T
u-e pbying thor put in company with th~ atttn&1
H

n($e m~
~intenscs[UJYJb.uto~I~·1S .'"'5
.{ th
:;. e ~~Ul-
f.acrors in moulding the. anim~ through th~ uccessivt
r:a.ges of ontogeny_ \\ e now W1nt to .... hQ.... .
· '.!'lCr c::nnsmiSSJOD tram ~t to om nng. part is plar<d,
l;ol <ll " deli f.t
'I'k'f~no",ctlkd ~Ien . J.O. ctors.or gen~:in This question is only beginning be ~ ed. anJ
~eJ 1:5 diso-ete Units SItu2ted In or on those the lines on which it is to be: 1.IlS"uN b.~ bttn
~ •
-- - "roen . the nuclei of cdls-the indiOlted by Goldschmid. In his e dettt-
_ es. A 1nge in cxd in one these min:ltion of sex in the gypsy m 1 he ius ~ tlat
e$ • - a.l " UD.OOn:llld the gene i then tixai the structure 0 the :idult) :is to weer it is ak or
:-In is ~ , condition until it mut2tes 3g.tin. It fem:t.le, depen s on the rd-J.tin~ s~ 'It which
is DO'" usutl regud these mutations 2S responsible sets of enes produce their effects. Th~ se
tbt'1.pponnce novel'~ in evolution. may be rd'erred as the m:olo-produ ' , and e
The ny in ",b.i e gwcs :ue sorted oU[ :Uld fem:lle-producing ~15. In other gm.es ss
Jistri ",ed bet.= puent 2nd offi>pring is "dl
~forgua's continu3t10n - ~len­
f.l rs do not only h:l'~ qU:llir:ltn"'e ues, etch... e
'Cll2S1. res: t :uso has 2. specific qUJ.ntit'ativ~ ,~ e.d roetion-
~ 's bJ, ana iorms the subject-matter the saen e rue. Yurious nees of the gypsy moth h;l\'T m"lle-
-~etics. Ln e pro«:ss . their tr2.nsmission the producin (l.l1d fen,:oIo-produoinJ gues 'di,l""nl
• es 1re curied in e gttm-cell the egg 21ld the qU:lntit3ti~ \Alue. By introducin_ in an __ con-
Sf"""
:. . .
G<rm-celJs,
. r e 111 cell' only mse by the binin 3. w feml.le-producing _ ne I sptrCl
QTlSlOn ~ on. dl :md the produ tion of nbining 3. strong nl"1le-producin ... e (15 CU! ~
n~ gtrnxdls OttUpies cdl-gener.ttions. It is done b' c.rossin moths oi dilferent n~ It IS
· ~ esc NO ~nttl '00- (which culminue possible to con\"eft \ uld-be fenl1.le:s into mlles more
'" Icniliu' ) >t e distribution of stenes ttkes or less comp1etdr. The con\'t:rSion es pbce It
~ te SO e gene .cist who s dies this distribu- :i cert:lin rime in e\ dopmrnt when the :;
' -.I (th fern:ole-
t."l c:ro:ituy tnnsmission is rallyHowing the male-produ in gene o\~.t.kes ,Its n\ e·
, •• ' . t the len\lle-pro-
es.. ...... two cdl-gener:uions only. But it t:1kes prodUCing gene), p to wIS PO" ,
• _L. d d the ~m".1 hOlS
-SIX ~ 'r,s cells to produce a body like dUCIng gene h:lS ~n 1l1le:l an '.L.

Q! " l:n:1n ou of "I'

~ ~I r .
been de,-e1oping :tIon lenllle tn""""J.
. . . but trom we
•
2.40 ~m' :i fertl ~ud egg (itself 3. sin Ie
It.
:01- __4uCI02 gue IS
.... ~ fifty-SlX enU2 'ons the enes point of con\-ersion on, the m~""" -
 •
•
• •
-- ~
.. • Ai

•
• •
"

1"" •

• •
:J%:S: -

•
- IUt
•
•
.
:-:'",":";, Y" • J. •
• -i;
"

. "
•

-. • •

-. •
•

-- •
•

-•
•
! . "

eua:s ..
--
...,. -
•
 SPEEDS OF DEVELOP.MENT SPEEDS Of' DEVEL t'~It~:"O l ••
o

"
bls eres
0
In G amnurus. ThoIS condu_ nte5 at which the thyroid gland de\'c:lops 111
mod> anf dconsider:able mterest,
0 "0
or It ena hi ('5 us to
5100
o 0

150
k PInce. .I t may be
• hNC'rochron)' aUt tn"C
st"e h0\\ . I dO
• • _oJ that a char.teter II was e\'o "e 111 ph r10geny
ml3.~nru .
btfort • ch:ancrer Bt but there 15 no reason why in
c.hm ter B should not arise before) simul_
onrogto r
ranrous!r with. or llftcr, h:u-acter A) according to
rhr rt!1fi,'c spttds of the gent's nt work, :lIld the
C'n\1ronmennl conditions. 1\ charncrer which ap-
pemd in the adult stage of the :ancestor might
de\"tJop elrly in the ontogeny of the descendant) nnd
,.ice ,'ersa.
In the verrebnlte5 the development of se\'eral
structures is under the concro) of chemical subsrances)
the hormones, which llrc produced in speci:t.I glands
2nd circubte in the blood. In p:uricular the thyroid
hormone is of interest here, for it is concerned in (he
de:\'~lopmenl of Ihe limbs, the lungs, and the longue,
i.e:, in the prod.uC"tion of those struclurts which will
turn [he tadpole inlo (he frog and so bring aboul
the c~!lnge which is called met:unorphosis. If extra
thrrOid hormone: is administered to n tadpole it will
m~t.lm~rph,ose too SOOn i if its thrroid gland is re-
mo\'ed n ",11 nor mc:mmorphose at all and the limbs,
lun,SSI and tongue nre not developed, The lime at
whIch metamo h . , .
d'rr rp 051S normally occurs vanes In
'hInt:renl
d'a
frogs llnd t
03 s, an
d H uxley has shown t h9.1
e lnt:re.nces' dol
o I In e\e opment of the followmg
each: .
Bufo lentiginosus (tOo'1d) metamorphoses In earl}'
summer.
R:IOi.l temporaria (common frog) met::unorphosc:s in
late summer.
.
Rana c1amirons meta.morphoses in second SC::lson.
Rana catesbepmn (bull-frog) metamorphoses in third
season.
The Ul}'roid gland de\'c:1ops fastest in the first, which
metamorphoses while it is still of mall siu, and
slowe-st in rhe Inst, which reaches a much larger size
before it metamorphoses.
Now there is little doubt that the sp«d at \yhich
Ihe th)TOid glnnd denlops is itself under th~ comrol
of genes) for Riddle working- on the thnoiJ of
• •
pigt'Ons came to the conclusion th:n the ;normal
I~le-chanism of he-redity can ope.f'3te in th perpetua-
tion of s.mall change'S in the endocrine organs'. The
~ifren'nc('s bet\\'een the to~ds lind fro .. JUSt men-
tioned mn therefore be indirectly ref('fr("{i to the
diR'l'T('nt nues of action the gene~_
, In the mammals the horm ncsllcquire-an incre-as('({
Imporlance in r't"gubring the sree.i of d('\'elopmelll
and deficiency in the thyroid f, r inshulce is weli
kl\o\\:,~ [0 be !ISS iated with the under~e\"t'lop('d
COndition known liS ulinislll. \Vhen th b I
of th d'c- e :I lnCe
0

e II1('r('1\( h rmones is upset the f

amma S CUn be I d de\'el ) It"mpo
_ °rment Ill!!,' be Ycr ' s('riousl • nfl~c, I
0

exp \une On the nsis of the diffe-cenl

e\ as tn
0

'OM '"
•
 ~r&I>"'~
SI'RP.I)S OF DI'.VRLOPMFNT 17
z6 .' d cribed by Gilford as Progeria ill
th c: Condluon became
es . an d
sentle id
extreme '
Y ccrepit rca II y , hc tadpole 'lage of a newt.
(AmblystOlua)
' lIonti'
which a ~an h had lived only seventeen years. J'05sesses gil1-slits and uternal gills. 1n ttll't state It
bY the: edlime:ith other
c: h d f
mammals, t e spee 0 develop-
can bccomesexually mature and 10 lOme feature' of It
structure luch :10, the bonts of the kull ne\ltr become
Com~ W 's very slow, a.5 can be shown· in the
mc:nt In man I • d b IJo properly de\leloped..Another ~n~ple j, ~~e worm
. mpariso n which was ma e y Ik:
folloWing co I • Polystomum integerTlmum, which." p:t.ra ItlC on the
monlh, to reach 3'5 kg. weight, and
Man tJ k" 9 frog, usually in its bladder, whe~e .It tak« three yean
doubles it in 180 days. to reach maturity. Hut should It Infect an early tad-
Ox takes 9 months 10 reach 40 kg. weight, and pole stage of the frog, it remains in the gill-c.ha~I?c:r
doubles it in 47 days. and becomes sexually mature in five weeks. 1 he
Some: animals go on developing throughout their structure of this neotenouS form differs from that
Jife, but most cease altering their shape at a certain of the normal worm in that it has but one male repro~
stage in ontogeny which is called the adult, and which ducti\le gland instead of several, and ~ther structurell
is characterized by the fact that the reproductive such a5 the intromittent organ, vagllla, and uteru'
org:lns Irc then ripe and rcady to propagate the race. are vestigial or absent. It is interesting to note .that
Now, the time at which the adult stage is rcached is some animals have become permanently committed
also governed by the rate of action of the genes, to this neotenous state~ so the newt' Proteu 1 Nec·
~ithcr directly as in the gypsy moth, or indirectly turus, and T)'phlomolge rl.":SCmble the axolotl, and
by metns of hormones as in the case of frogs and Polystomum ocellatum rC'stmhles the nootenou
newts. It is possible, therefore, for there to be a Polystomum integerrimum.
competition between the genes which control any On the other hand, by delaying the time at which
particular character and those which determine the the adult state is normally reached, it is possible to
assumption of the adult stage, and unless the former cause structures 10 appear which would not ordinarily
w?rk fut enough and get ill ill time, the character have done so. So in the case of the female gypsy
""Ill not ~e able to show itself. This is what actually moth, Goldschmidt was able to delay the onset of
ppen, 1n those cases in which an animal becomes maturity sufficiently long for the male-producing
xually mature h·1 ·11·
w Ie stl In the young stage a genes to overtake the female-producing ones and so
henomenon kn '
..ow n as IIeoltll] or paedogellesis. An bring about the formation of male structures in an
I
xampeofthlSlsf . hdb· .
urllls e y the axolotl, which IS otherwise normal female. I t will be shown later that
 OF DEVELOPMENT
SPEE DS
])HYLOGENY
2~ . like this has possibly taken place in the
sOl1le~,"gof certain kinds of animals. perfect, and we only have a
,
b f
• num er a more or less
'9
c\·olutlOIl.t. then we may safely conclude that the Isolated forms to indicate the ",ck h· h h
W Ie p rlogeny
AltogeUJerJ J k f has taken. Nevertheless, the studl' of the h I ·
. h" h the internal factors wor are 0 great ' P }' ogehles
needs 3.t W Ie d .. . f b
o anum er of ammals such as the horse, e1ep hant,
Sr· . development, an variations In the ,
importance I n .
.
.
d of the vanoUS factors may play an lm- ~nd camel, have !1e1ded very interesting series show-
relaove spec 5 Ing the progressive modifications which these races
in the relation of ontogeny to phylogeny.
portan t par t have undergone during cvolutiol1. Now, on COI11-
!)aring the various members of a phylogenetic series,
IV It appears that the differences between the structures
PHYLOGENY of the adults arc largely differences of proportion
and of number of the parts relatively to one another,
HYLOGENY is the rehabilitated scaleofbeingsJ
P a row (or rather a number of rows) of adult forms
which are related to one another, not from adult to
D'Arcy Thompson sa)"s in this (annexion that 'it
is certain that, in particular cases, the evolution
of a race has actually involved gradual increase
adult, but from the fertilized egg which gives risc or decrease in some one or marc numerical factors,
to one adult to the fertilized egg which produces the magnitude itself included-that is to say, increase or
next. The adult forms in phylogeny arc therefore decrease in some one or more of the actual and rela-
disconnected, save for the (unproven) possibility tive velocities of growth', In other words, phylogeny
mentioned at the end of Chapter II that the external in these cases has been brought about by variations
factors which control the formation of an adult might in the relative rates of action of the genes, which
become converted into internal factors > and so affect formed the subject of the last chapter.
the ontogeny and production of the adult of the next The comparison of one adult form with another
generation. can be made very instructive by inscribing the shape
, of one form on a grid-system of Cartesian co-ordil~­
, If all animals had been preserved as fossils, and
If ,II these f·\ . h ad been discovered
. ates. If the skull of Hyracotherium (an early fos~11
OSSI remains we
should be able, . . > 1 regarded as ancestral to the horse) be in~cribed In
a trace a ContlOuous senes of adu t
ancestral form h' h this way and the drawing be compared with ~hat ~f
s W IC would represent the phylogeny
o f the race wh'ch ' the skull of the horse, it is found that the, points In
th' , I we were studying. Unfortunately the horse's skull corresponding to those 111 that of
IS IS not the ca
"
£ hi' , '
se, or t e 10SSl1 record IS very 1111-
 PHYLOGEKY
PHYLOGESY
F . b1\"'C been. displac~. B ~ a harmo- efully d . J'
ue peac . un ergomg qua.ntitatin chan
H!l~ Don of the grid, the skull of H}-n- uch an evoluuon of structUres' th . <So
.;.us ~ distorted and "",de to resemble that e J'OUng IS ",ell
(X)lbcr.'==But:
o ere are in _0._ e s cps m- th-IS
alDCUl-"I
kn own, and called cae:nogeoesi<
.
UI
...., or }"'OU thful adapa
non. If now neoteny OCCUrs, and the aninuls -
of t1l< bar><-znd "" :aauoJIy h..-e fossil horses which
sexually mature in the roung condition, e brlo-
t~~:a:xCi2te cPs perfectly (. esohippus geny will undergo an unexpectedly abrupt ~6c.-
~ Protnhippus). We ""'y wee Wlth D'Arcy nOD, and start off In a new dirtttion atrog th ,-
il c et. -"ow ,
'f'ho,-17"' nen he writes: . I ~ IS ~e ratio ~tween
.L ,

~ n;es c1 groonh in -nrious direcoons by which we
_ account for the enernal form of aU, save certain
~ minute, organisms' j and we mar conclude
~ du! it is an aJ eracion in this ratio between the
r=sci g,ooth ...hich h2s produ=l a phylogeny such
.. tha: of the horse. It is DOt n = r y to imply that
...Jlo.=ioc bs ocly consisrrrl in quantitative variation
t1 Uc:aCy caNing strueturcs; on the contrary, quali-
",,-.. n..~ in e form of structural novdties
"::~zloo h:i<e appeared_ But one<: the internal
f: f« dtese DOveities have become established,
they will be subject to qtwttibtive mcxli6cacion and
'-rochrooy.
It is, perhaps, -.rorth essing the fact that if a
~ 'Ppeal"; and only affected the roung stages
. .-ag0l! 1D a ntt, that race would nOt show an)·
FJIogerroc progression, sine<: that is measured onl~
01' ph l!1Odi£" J qU2licati"-e novelties? In the following pages
.• . camo. I IS therefore possible to
t:=agt:e d:ar a <=ain ._.- f 'cl d . •
Doe. of ~ ~ 0 an esnne ~olu-
_ ....oJ es may take place in the
""gts or development while the adult stages
. record
LUe lOSS
.has been of the grenes _. ~~ue 10
.
tracrng the phylogenies of horses within the mam-
malsJ and of mammals and reptiles in general within
me ~eneb~ ~ ~ orJ to take another group, of am-
monites WIthin the molluscs. Bu no fossils ha,,"e
hoen found which bridge the gaps between the I er
groups (or phyla) ofanimals; bet.-ec:n the Vette 1eS
on the one hand and the starfish on me other 4X'
•
betw"een either of these and the mol1usc:s or 1rot
Doubtless thae were many forms whose fossil re-
mains have not been found; there must have htta
still more which were not fossilized because they WeI
not have an)" hard par capable of being pres<n-ed.
But even so, is it not possible that these gaps, that
these discontinuities in the phylogenetic series
adults, may be also to a certain ex ent due
'clandestine' e1o"'Olution in the young sugcs folia::
by neoteny and the sudden revdacion of these hid
attempt will be made to show that this has oo;un..
in some cases at least.
At ention nay now be paid to those cases
 Pttl ........ ~-
0: .' h the modifications of successive PHYLOGENY • 13
J •
brlnuenJe5 n ",hie . taken place in d e 6'rute d'lfec- constant dependent on the particular case,and"leiS
P "-:,0
~ulu appear.
I
(0 ha"c .
a constant
"
which IS usually about 1·5)
int is dlat of the (ltanotheres, a . . Th.· . . IfitO'~t
cions. A esse .1Jl:O mammals. The earliest fossil of
. thiS from
. the. present point of view,', ,h at any
group of ~ borns on their heads. Later IOcrease tn ~e ~lze of th.e 1:xxl.y will necessarily entail
ri[7.lJOtherts b nOd in each of the four races into an exponenttallOcrease I.I1 the size of the heterogonic
il had borns, aD
foss s _ th es seem (0 have split, these horns or~n as a ~orrdated variation, brought about by
· h the otaDO er .. an Increase lh the Tate of action of the factor con-
wh!< . I' larger and larger, unnl they
e rogrcssn-C) trolling. the ~ormat_ion of the heterogonic organ.
bec= P'dieuJ dimensions and were probably
Complying With thIS, Osborn has shown that it is
r""bed ed'w
n oUS
he extinction of the race w h'1C h t h en
concern 1IO t od a fact that the bodr-siu of the titanotheres did
This incorrigible tendqtcy to pr uce
ed
",curr . . Iy 'In t he increase, and that in those forms in which the horns
-horns
larger an d Iargo.. ,
to vary connnuous
were small they only appeared in the adult, whereas
same d·IreCtlon,
. has been given the name of orlho- . in the later-evolved forms in which the horns were
. While manY authors have attempted to assign large they were already present in young stages.
Iuum. I . .
orthogenesis to the action of myster~ous forces, us Further, it may be concluded that unless the size of
probable explanation has been furnished by J. S. the body has reached a certain minimum value, the
Huxley, from a study of the relative sizes of parts of heterogonic organ will not show itself at all. So while
animals at different absolute sizes. In the case of the the early titanotheres were too small to show horns
anders of deer, the ratio between the size of the antler they transmitted the factors for horn-production to
and thesiuof the body increases with increasing body the four races to which they gave rise, in each of
size, or in other words, the antler grows relatively which horns appeared independently when the body
faster than the body. Many other instances may be became large enough. Orthogenesis of a strucrure
given of the same phenomenon which is called in phylogeny may then be regarde~ as due. to the
htta-Ggo,,}. In all these cases the relative growth-rates existence of a particular mathematical relat:t~n be-
of the heterogonic organ (in the case mentioned, the tween the rate of action of the factors controllmg the
ant~er) and of the body remain constant during long production of that structure and the rate of action of
penod,: aod may be expressed mathematically. (The the factors controlling the size of the body. Lastlr,
~resslon lS-J = h,t, where x is the size of the it may be noticed that if an animal with a heterogonic
· 0 f't
organ were to decrease t he Size 1 sbo dy, that or"'o
y, J the size of the heterogonic organ, b is a 0-
".. ,
 IIJ','I'IOHKIII\ONY ANI) J'IIYI,IHdNV
II
11(' lull nHllr ~IHllplh_.1I d 111 VUhlllCIIl II,
ry 1I1(1~,
IOIlW" III dry lUl'l whh.h rllnnl lh I III tu,
• If" t
whh.h Ih rOUIl~ IlIllnhll w" h lhhnl nd IliuM I'Ml
IrlldiliM fIll hllltll w.. lit I.l} d. A II 1I.llll"ltllll II'
Ihl" \!f·h1Y n "fill of flllHI IN.. I'rnvlllni fur II !.-1I1l
·""Jlllioll hy Ihr IIhllh. I in Ih.. form ul )c)l\r. 1/\ Ihr
'~H' lSy ,hi" 1Il1'.11l IIw tllllll)Clllh p("llu~1 \\.l pm
V
IlJllWd III !Ill' c'xpc'u1r 01 Ih4'" I.llv"l. 141\11 HIiIIUf('
III' 11'IWtl/llIN\ ANI) I'IS I'."",", "I' whh h hili-! 11 'Ill ItltVlll in PI'("VIIJlI 11111111("111(' .... lHll~1
IN l'IIl'I,O(:",N (111\1 111M IIH'y Wfl,·d hlyrd!ll 0) CIlmr III hrC'llIhr)'UIlIL
I', Ill\' rfll ill Pll'yjflll~ I !lupll·r 'hilI Ih l
Whrll,l!wll'llll(', it itt dl''1i'c-i111l pC·,Ik. !lIlA IIIHl\ur

W "'lrllu1hfll III Ihtl IIlltllhd 1111101_ 01 d('vI'lop

11lf'1l!41WVlllY~lltdr.rll !lII'1I lil"11 ~d dlfl(1rrrlllltl(',.
01' (hnllH Ir,' whit h llppr.ll'. in r,l' I)' LtHllnllln-dtlll
tn( III It 11l.IY Ill' I,ll hrr IHlll )'UlIH til' hu VI\1. Whll h il
.l(llllllty lit Il'I('1i nil! IHutlC'1 11)1' tlw 11I1'lirlll plUpn r.,
willi Illn 11''f1l111lwl dl~ tllllf1 01 Upprrlrli,lt flj'U tilliH
,Inil I), In oldl'l' 10 11\'11111 III\lkll\~ ~III\Ntilllt;ilti WIWH'
lut HiliIt ~llf rrd. 11111 dill" plJI~~lhl IlIl' IWIlIIII.(lll\tt
llljllf' j/4 l1lt'~ltll, III 11)(" lollnwlllH p,ll-\f th~ Wlllll
Iq If'WIII ,It 1I111n /II 11It'11 IIPIH ,Illilli (~ Ii Nut ('_1"\,(1
, '0\111,." \\1111 hi 1I/4('d III I I'll lit n:I'H'tj 1111 I ,Illy l.l~t'
olllnijrllit I lid, hy \I~lIyjllH Ih I ~11f" III wldl h llnJIlHtI"
01 Ilf\vt.lIIPI,l(·"I, 11111 Ilw Jill til 1111 C' \\ hi, II ,Ipprolr I
hrilull l1\lIlIUI, Jldlill lrullIlIl' (.111 lit' I(·dll ,'d 101
,''" hlH will h \.111 d' lHllhl .. l'.
II \lr~ljN ItlUlllj i.lllll·d,IH 1IIIIIIllli 1111(1111 111111\ '11w dl dill dOll lit'l\\' 1"1\ Ilduh 011111 IHlllj.l" 1,('.
hrllllllr. lid II II ~111I1 II 1If1 1lllllHlw d jilIn Ih~ ph dq hrl\\I('c'll IIllll,llllf \\'hllh 111'1'1',11" 1.11 III t,ld , 14
wily. A w"rlllllll,1 II .tId filioli' Ih II fill IIillldlll, dltlWIl pllli IPfl II IU"lltl C' I I I I ,nl' Ilw Illlll\ltlrr
l'f'UI! Ihl\ljlllr llir I~H I' Inlllllr" 1111111 ,II nUll i,1 III \lllllil \vlllih ~llr 1111111'11\(,1 III ph IIIWIl.

IItll . rlllnw flllllllhf'l fHH IlIrll II II ~lIlc ,I, • hllfl Ii ",

,ll tl
Nfl\l,f, ,Iii' PlllHlpl fir It,'rli\,hllll\ Will IH,Ikr ~l
It I. kllllWII ~ II I IUhr)lI l \Iud Ihi pi I lid I t 1,11t'1! 1'1I.,t1lJlrl Iill 1111 1I ..,l1lr III \
~'1I'C'
'1 htlrt III (',lIh( I
'"11" ,uli, • III III Illlh IHII • \.llld unlll I hu Il~l'illlllr. I \I well ~IIH" 1111 11111(11 011 ( 1'11I1i I 1I1~
I Ill' "hCII 1,)11\
II ,
Ill", udult loll 1 I /11'1,1 Ill" 1111 to
ll\tllj.\ 1I11111Wl I ~ll/jWIl ~lIi II Ihll('d wllh II I"l'\ltllli
I. I I, II ollln/1I'II, ,
Ilf 1'1l~J11
""", 'hi. I",j ,I I I II 'O~llhl\llllnd III ~1l11111 "llllllllr ",
WII II I
n I"'
II I " " uH "I I tI "" VIII. \VIllI< IUIH"' )r
J....l' )1' I 1'111111 )'lllIlt
_ Jlf1dqd W~I" y I hlld lUI IH1Inllll" hllld., 1\ dl,b, I'd 11PIH'111l1lHCI I
IlI'\\ 1111\ Ill,
 RO<':M~V'" ..•. -
6 HETE "I
3 . duction to vestiges un e55 the time HETEROCHRONY AND PHYLOGENY
"II can their re d" I
WI ~ h dult stage is rcache IS a so delayed· on all the internal factors and wI"11 t h ' 31
t which tea .. , LastIy, a d15tJnCtl0l1
"'" f' , n o t erelOre be
:l , th possibIlity. o s.uch Importance as the internal factors in ro-
hence a se,en hful
W d between those rout
ust be rna e d
structures ~ucmg heterochrony. In the following consilra_
rn . only the young stage an those which tlon.s the external factors are regarded as constant
whICh concern
. d also affect the adult. Heterochrony, dur.m~ the. phylogeny; the possible effects of their
"",Slst an
r-- r rovides eight POSSI"b"l" " 0 f vanatlOn
I HIes "" in vanatlon Will be considered later.
re
therelo , P .
the way in which structures may appear 10 an onto- A ~haracter which is present or makes its appear-
ance m the young stage of an ancestral animal rna}'
genyas compared with the ontogeny of the ancestOr . in the ontogeny of a descendant appear:
It is of course impossible to measure the relative
A. In the young stage only, producing youthful
rates ofaction of the factors in controlling the develop-
adaptations or caenogenesis, and not affecting
ment of structures in the ontogenies of all animals,
phylogeny"
especially those which are extinct and known only B. In the young and adult stage, producing a sub-
by fossil remains. However, there is the method of stitution of a new adult condition for the
comparison, for if a structure is found in the adult old, resulting in progressive deviation in the
stage of one animal and in the young stage of an- ontogeny of the descendant from that of
other, it is obvious that the structure has developed the ancestor.
faster in the latter. If, further, the former animal C. In the adult, by a relative retardation of the
is for other reasons regarded as a representative of development of the bodily structures as
the ancestral type from which the latter animal compared with the reproductive organs,
descended, then it may be concluded that the phylo- resulting in paedogtnesis and neoteny.
geny of the latter animal has been brought about by A character which is present in the young and adult
a r.elatlve acceleration in ontogeny of the rate of stages of an ancestral animal may in the ontogeny
actIOn of the .factor controlling the development of of a descendant appear:
the ~tructure 10 question. All the eight possibilities D. In the young stage only, resultin~ in the
"0f
Outhned above may be tested"In t h"IS way. It IS reduction of the character to a vestige.
Course true that th I A character which is present or makes its appearance
. '. e externa factors can also operate
In moMy,ng th f" in the adult stage of an ancestor may in the ontogeny
. e rates 0 action of the internal factors,
but they will act h of a descendant appear:
On t e whole more or less equally
 , f'IIIUIN\' ANII 1'1IVI,(H)J':NV II wn:ItOCIl ItONY ANI) PIIYI , O('!'''Y
1l1,'IIHI • "" :\9
" I I 111I1o'u(' Il'lIldtif l K III lho r tll/rn('lhc~ hy t~l 11.1(' d bu :11110 by the magnitude of the
I' 11I1,rl4,II ~I " ••
willi h d,..'lllJ.pllllh 1lllllVidu,dlll, VIlI'lr(1 '1.llI\d rC,I~II(}Il) I,e. h~ the amount of agent whi(,;.h i!\ re.
'.11".: 1I,1t,11 't'", I"tll/H, :t ling, :loti till may ;1150 affe t the aile f the
I, 111 fhco \",r IUIIiIi ..t.l),lr, i. . Inn 1,llr, I (,'Jllltill~ lruc~"rc. It .apt' ,l.ra, t~crcforc, to be necc:, :IT)' to
'" fh~ tnhl\ IIIUI ill Ihe' I hllllllirf "1 a vcMI if.lt t,lk ,11:tu toll 10er,1.1101I d~fTcr.c:ntc of c.:ar:u;ity a, well
II, ,,/./,.I,t" 'Of. a dlflc:rcnlc of mtell'ilty In the rca ti n of the:
t,. In lhl' Ill! '''N('t whith i IlU InrlH I •• dull, Il\lcrn;11 Eu:.tor!'l in producing !\trll(ture~. This i~
the' 11r\V IHI\lIlIlI.I~r hrill'" I ('1.Hively drl,l 1('«.1. ~howl\ hy the work f F rd and iiuxle)', who ~ und
If 1l1"11~ III '11\'(1 lrPl'illj.l' ,11(,' I" cviou'l Ih.llth condition in which Ihe C)·e: fGamnuru· i~
\ IIi" IN<'IIlf (11 If r/,f""'" /,llo,li ~ .
(l.de (i, '. (Onl.lill~ rdativcly lillie hl.u,k colourin~
1ll.lll r pl'r unit are;l) call he ohuincd not onh hy
II III lhr IHlIlM hl),(l', plodudllJ-l, IUI-'(fHiOli
.l dCll·C.I'icd r.Ut' of dcpo~ilion f the hbck (i.t,. h)·
I'pprJI,lIl1 III tht' .lIlH'·HI,d th,ll,lllt'l' .1IH..1
dnl.t"l~('d illtcn,ily) but .lls hy all inl1'e:al.;c in the
""d".lhlJN. ,Ire,l ofd\(' t'ye (i.c. ;n~ullicicl\t cotpacity). (onvcr"cl}'l
t',l1l' 1\ lIHd • whidl prod\lU' ph) 11lf.\l'nrlk cll'c:u'J
ill (W\('~ ill which til(' .In:,1 of lht' C)'l' is snull (a.. in
h, 11lIt,\dll\1tl~ \\l\llldull..h~l\.Hlt'l' inti) lilt' lint' flf
dW,ll{$) \I ~t'lI(, of low intcnsit)' h,IS sufliticllt Clp:llil)
Ihlull Ill.' I 11.. . \\\illhilln\ lltltk,' thl' {l'l'lll fliI(dulmJ"" 10 ~dlO\v .1 ridl depo~it of hbck colourin~ Tll:tltcr l ;I..
f III, r,l~r F, C: l .llId 11 \\ hi . . 11 p1'l)(.lllU' ph -hi
W,I., !,,·o\l\.'d h F\1rd.
'IIlrll, UrI! 1\\ llmdll' inJ.t "h,II"111,;('1' whil..h W('I'(' III S,01l\C (,\.. e~ tht' time (If .lppe.lr,lI\lC of.\ ch.lf.Il:H'r
:l1'fld\ pIt' (Ill III thr 1\ll('\lt',11.!1l11 1ll.1 . l1\fcl1icl1tl m.\ be lit-I.I C\\ h . th ;\Ction of :molher f.Ktor whid\
hI.' lIId\ldr\\ \llldn Ih(' (rflll ,~" HI rNlJ,pIW.I; , .Ill" .IS.t (\.'mpor.lf)' inhibitor. The fenl.tlc-proJw.:~llg
11\ Ihl' 11 111,\\\ II\~ l'-'~( I (' -,mpk ,,111 h(' i\'clI tlf ~cnc'" ltCI .1'\ ec.::l1lp()r,lr) inhibitors to the prQl,.hl(~lon
r.hhl1llhC'(' It \.\. ,
I \ 11:1 \ I~\""I II IliC l.1\Ithll d ,Ih(wc. J\ttcu" of llulc ·h.\r,l(lcrS in the g~ p~ moth up to the: tI1ue:
h 1\ 1111\, lhl\\f\o 1\
\ I \ C \,t f,
1\ I 1\CH' 11\ .\ l'mllt
" \d""h .It \\ hid\ the l1\.1IC'-l'rodul'illg' f,l(lor.; win .lId the
1 Il\('lt' th(" 1111 '
.1IliIlU! is ::-witchcJ vcr from nc st" to th(' other. .
1\.1 1 1 . Ilncr III "Ill"h hr(('l""hrtlll 11\,"
\ \\ (II \hlll~l\t \\h I 1 I . h' I
1\11(" I' I (III, I It '(' II u{" th,lt .1 ~I' ur It pocs, of l:t)ur$C, wilhoUI s.l)ing th:lt t c tnllC ()
\ nn('\ ~I "
I \rl I.1111
' , .lppe.lt.lTKC 01'.\ dur.IClcr 11\11"t he subsequent fO lhe
"I ~ \fll.\' 11111('.\ .. 1 ( .. \lIt \,1 .1 I (.h..li''li
, I' I I ' IroH'ln r,. it Ius bcC'n set
1I\1\,1"t(' lit" III'~c\
I " thCl\ I II tl \(" 1''''('~Il~'11\\1'1.'.'" d ,h\.' tllllC .Il WII'" 1 I lC rC:ICtlOIl C 1\
. . .
1:\
. ' If bcillll l'OntrolkJ h)
I ,(' l,lIme I (' I, II ' '
t\1'1\('lI;ltl\'~\)111 __ \ .U U:I", \II f!l(' tnllC \)1 III 1ll\)f10ll J IlllS l.ltter tlln Ibe: 0

1('''lllltl\Il~'llI\1 11\' \.'lV' \(' II

 o Jlg'J'I~IH') /l1(VI" ,.- • CAI~NOGENESIS
4 . which in lurn 18 governed by olhers
'. .'
t he ear Iy stages ,of ontogeny . It is also worth nOhclng
rO flO Il
~n().ther ( Ihe mottlent r the lerllr 'I"Izatlon of the'
th a t th e evo Iuhon of the mammals co ld t h
1 . I1 ntogcny IJCgIIlS. '
a~e
".fnutt r01l
I' ", I ',the lillie at Whle
(Ul!WhH.11 Ih
taken .place if• it had not been for thes"e e m nOb tYOntC
rr' II I"" be undU'1 d lhat a l ollgh we .peak novelties. It 15 equally certain that the Glochidium
It WI a~ .
' tert :tppc:lrillg c~rlr r late In nt geny, we larva of the freshwater mussel (Unio) which is tem-
II r l h,If'" ".,y file InS imply t IlIlt t Iley 3rc .lin'It char. porarily parasitic on a fish; that the larval stage of
do 1101. I'Y'"
" the s Ie e(fects f the •Illternal fa t fS Haemocera which is parasitic in a worm while the
Quer , I. .
onlrollinf{ ,hC'l1l· adult sta~e is free and resembles normal Copepods;
that the InAated larva of Sitaris which floats on the
VI
honey of the bee Anthophora which it parasitiz.es
AE E ESIS while the adult is a normal cantharid beetle; that all
I1ERElirc a number f :l cs ill \'/hieh differellt these youthful adaptations were introduced into early

T !\lIim,l1s resemble II :\llothcr when adult but

!\rc Ill.trkcdly unlike one :1I1othcr in til' :uly stages
stages of ontogeny and reAect no adult characters
at all.
The Nauplius is a larval form commonly found
or their dcvcloJl1l1C1lt. me f the char:lctcrs which
in the Crustacea; the trochospherc is a larval form
thesc ltn;rnn!s h:lve in their c:wly stngcs could 1I0t
of frequent occurrence ill. Annelida and Mollusca.
pO!isibly 1I,lve heen present in the adult st:1gc f :lIlY These forms must be regarded as larval adaptations
:.nccstor1 and so it is cerLlin that they ar sc during to dispersal, and not, as recapitulationists contend,
r.1r1r 5l.lg( (development. This (:lct is important) as the representatives of ancestral adult forms. The
for It pro\,( th'lt cv lution:lry n vclties cnn :\Ild do trochosphcre could not be the form of the adult
'pl'.,r in mly tJgrs f ntogeny, Here belong the ancestor of Annelida and Mollusca, nor could the
(~ f ('mbr oni and 1:uwl ndnptnti 11 to which rauplius represent the form of the adult ancestor of
11:1(' kcl :lpplied the term {.u1Iogt1ltlicJ n few of which Crustacea for that ancestor must have h:d it~ bod)'
J
'1',11 1nOw be 00' n 'J I ertlI b W:lY of 'Illustrauon,
' divided into several segments, while the Naup.hus has
.No ne: WII\ deny Ih:\t the. amnion and the ::tllan- Th t m any Annelida and
only three segments. a so .
10'S (embr"• • 1\1 ,
11lCmOf:U1ts)J which arc of such Mollusca do have a trochosphere larva is d~e t~ ~e
t'SS(1l11l11 IInl rt:\
,1\ e t
I
t le dcvd pment of every fact that the)' have a common ~lI1ccstor (whIch tIlth"S
'\ I,'
t') ulrd ) nd ma11lllU IJ could never have featured
I ,v.t) whatever at
r<'pll
• ontogeny had a tro ch osphere a
1Il nn ' adult anceSlfa \ r10rm nnd must h:wc evolved 'III lb6<l c
 CAENOGENESIS
CAENOGENESIS
.' h been like when adult. The same
may ave
ancestor '0
I'
I . n app les
the Nauplius. These cases are
.
cone US10 h t s the amnion and allantoIs because
trenC an a .
not SO . f argument albeit false, whereas the
th do pernttt 0 .
er. d lIantois leave no room for argument In
a111111011 an a • • • h
.' retation. A decIsive case IS) owever,
their mterp
'd d b the method of ce
11-<1'IVISlon
, , W h'Ieh the
pravl e Y •
'1'__ > gg undergoes In the development of cer-
fatl IZCU e .
rain Platyhelmia (flat-w?rms), ,:nn,ellda (true w?rn:s),
Mollusca, and Nemertma, which IS known as spiral
cleavage', In all these cases, the n~w cells ~re formed e a
by ctivision from the old cells In certam regular
positions which follow a definite rule. But this rule
cannot conceivably represent any adult feature of any
ancestor; it is merely a developmental device for
splitting up the single fertilized egg-cell into a larger
number of smaller cells, and for sorting out the raw
materials that will go to form the various regions of
the future embr}o.
Many other cases there are in which the younger
stages of development of different animals are dis-
similar while their adults are much more alike
th h . ,
o~g. It cannot always be asserted that the youthful
vanatlons are adaptive. Hurst drew attention to the FIC. I. The young and adu!t forms of thru IIio, IOS~OW the:m~~
ract that wh'l h d I ' between the adulu and the d.lfen:nas betW"tCD the )oung.. •
L' t e tea u t forms of the Insects Culex, b, Chironomusj (. COn:lhn. From Miall.
CIllfonomus and C h d ' '
J
1arva 1stages sh oret ra 0 not differ widely, their
very largely 011 the platform of the recapitu]ationists,
anoth S ow much greater differences from one , ' th em can be neglected.
er. path ha d . no eVidence concernmg .
of alf" s rawn attention to the same state Spa th sates ,h at ,'as ',n Illany other AmmoOltes,
airs In Am m . t
omtes, and as these animals figure
 CAENOGENESIS
H. dult of Gagaticeras are remarkably
whilst the:l p1:lstic young show great variability'
. 'bI' the more . . f th" " ""
sUll! I h r a ~i01ple application 0 e biogenetic
·Iti~deart 3 ' : ' 1 b
.) G3g:1tice ras would le2.d to resu ts as a surd as
~2W to O
f a new-born monker· To add to OUt
mtheost • there may b e pronounced
"
oyet'W e1m 11\
g detail while
h I
. . throughout several depressed whorls of
r:mcoScatlOn . It is dear, then, that evolutionary novelties do
r--"no>ticerasalready at a dlumeter of a few mm. I
roungv"S- . arise. in th~ early stages of development and ma.y
in other immarure specimens there IS not only fine
and close costation, but the whorls are round or even
compressed at sm~lIe~ ~iam~ters. . . . Thre ,larger
whorls are almost mdlstlngUishable. . . . \;\, hilst We
should not hesitlte to identify-specificall)'-all the
2dults, ontogenetic peculiarities, such as variations
in whorl.shape, might be adduced to justify the
assumption of the most diverse phylogenies and
skipping of h}l>otheticai stages.'
It may also be remembered that Sedgwick was
unable to establish any specific difference between
Periparus capensis and P. balfouri' except on the
structures of the earl)' stages of development. This
case is particularly instructive, for it shows that it is
possible for a certain amount of evolution to take
~ce .in the younger stages of development without
eetlng the structure of the adult stages and there-
fore withou.t producing any effect in )phylogeny.
Such evolution would then be 'clandestine'.
" h me of the products of caenogenesis. may be
h'gil' "I"Iud, as in the case of the Glochidiun_'
_ _ _ . SpeCI:!
CAENOGENESIS
larva of Unio) or the parasitic larvae of HOle 45
"d mocera
or Sitans) mentlone above. But as we shall
bter) th esc specla rz.e cases are unimportant f sc.
"I"d
"f " f rom
the POUlt 0 VIew 0 phylogeny, for in all proba.bility
the characters of such larvae cannot figure in the
adults of any descendants.
restnct their effects to those stages. It is equOll1)"
dear, as Morgan points out, that the internal factors
controlling the production of these novelties can be
and are transmitted like any other internal factors,
i.e. are genes. \Ve therefore reach the conclusion
that these youthful characters are prone to the effects
of heterochrony by acceleration or retardation of the
rates at which the internal factors act, and might
therefore be expected to appear in the adult Ols well
as in the youthful stages. This possibilit), will be
considered in the next chapter.
VI1
DEVIATIO
N this chapter we must consider those cases In
I which the young stages of development of different
th
animalsresembleoneano ermore an
th th.yresemble
the adult stages
the adult stages an d more than
I V h II therefore, ha"e fO
resemble one another. , e sa, pose his
deal with the cases which led VOIl B:ter!O pro
_
 nl:\ 1 Th
. ill "Itt--. r nthrr f th(' gill--
"
in th(' ('mhr~ oi.: t tt" tu~~~~
nul. I he (' ... rru tur ,
'""""ubi ,

•
 DEVIA'J'IUN
.. 8 5 sueh as the Eustachian tube, the
h r strudure d Th . .
or c. d he thymus glan s. ere IS simi larity
,onS11s, 30h t mbr)'os of hsh 311 d o f rcptl·1 es, birds
Ixrwecn r e e f '
15 but the later stages o ontogeny have
:lnd mamma J • they are adult .h~ve elght pairs of limbs on their
• _..I I tne reptiles, blrds, and mammals, other
diVerge<>· nhave been substltutc. d Jor
r he a d u It stage
t
,du It sragC'S
of ,he 6sh. During ,he phylogcny of the reptiles,
birds, and mammals, ~hercforc, faeto rs have arisen
in the ontogenies whlch control thc dcve,loprnen t
from tbc embryonic stage onwards and wh leh have
produced progressive deviation .
Tbe adult stages of hermit-crahs have curved
abdornrns while those of normal adult crabs are
symmetrical. Thc larval stages of hermit-erabs are
symmetrical, but this symmetry surelr represents
[har of the 10".'01 stages o f no rma I crabs and not that
of me ,dult crab. This symmctry of the larval stages
of larval hermit-crabs could on ly represent the sym-
metry of the adult stage of norma I crabs ( fro m which
hermit-crabs are regarded as descended) if phylogeny
play:ct any pare in ontogeny, and we have secn in
prevlOUS chapters thal thcre is nol s ufficient reason
to b~lieve that this ca n OCCur . Be this as it may, we
can In another case prove t hal a larva I form whlc . h
resembles ,he ;arvaI rlor m o f another animai
• .
. whlch
IS regarded
~ as ancestral, actua l1 y difl'ers fro m the
d
a Il lI IOrm of th
C al ancestral animaI. P o rtunton is a
rustaeean of th d
stage il is .e. o~ er l sopoda, 3nd in its adult
parasltl c In crabs . Thc larva! stage of
DEVIATION
. '9
Fortunion resembles the larvaof normal non-parasltlc
..
.
Isopoda,
.
and
'
hke the latter I it has sevenpalrso
. f
1Imb5 on lts thorax . B.ut tbe normal lsopoda when
thorax , and SO lt 15 definitely cIear tbat tbe larva of
Por~nion takes after .the larva of normal lsopoda
(whlCh are regarded as Its ancestors) and not after the
aduh. 'It is the adult Portunion which has lest iu
legs, not the young Portumon which ha, acquired
them from its adult ancestors.' (Garstang.)
Entoconcha is a member of tbc Gastropod order of
M olluscs and it is in its adult stage parasitic in 5e3-
eucumbers. The larval stage of Entoconeha i, a
Veliger similar to that of the larval stages of more
normal Gastropoda , and so here again we find thal
the larval stage is a reRection of the larval stage of
the aneestor and of nothing else. That Entoconcha
and normal Gastropoda should both ha\'e \'e1iger
larvae proves only that Entoconcha is a Gastro-
pod, just as a parallel line of reasoning proves that
P o rtu nion is an lsopod Crustacean,
The sole is a so-called 'Bat fish' because in its adult
state it has turn ed over on its side and lies Rnt on the
sea-bottom. [t :\150 undergoes a eertain amount of
struetural modification resulting in the migra~ion
o f the eye 0 11 th e underside on to the upper slde,
and eon sequent asymme tr ·lcaI d·s'ortion
1 . . But. the
larva o f Rat fish is symmetrical aod SWlms upnght.
and so is thc larva of normal fish . The symmetry of
36'6 H
 DEVIATION
;' f the Bat fish corresponds to that of the
the /arV2 0 I fish and only indirectly to that of the
,._~ of norma . h
..... f normal fish. Ir IS true t at normal fish
adultsage 0
. symmetrical while Rat fish become asym.
rem",'" I.10 the adult condition, and therefore the
memo
Bat fish bears a stronger resemblance to the
1UV2 0 f .
adult of normal fish than it bears to Its own adult.
But this is because the normal fish has not changed
whereas the Bat fish has, in substituting Its type of
adult structure for that of its ancestor.
The skull of the sharks is cartilaginous throughout
their life; the skull of higher vertebrates (e. g. mam-
mals) is cartilaginous only in the embryonic stage
and the cartilage is replaced by bone in the adult.
This cartilaginous skull of the mammalian embryo
resembles that of the embryo shark, not that of the
adult shark. The essence of this fact was correctly
discerned hy Reichert nearly a hundred years ago.
lD the same way, Antedon, a Crinoid which is free
and unattached in its adult state, passes through a
larval stage when it is attached by a stalk. Kormal
Crinoids are attached hy a stalk in their adult state,
.nd their larvae must be attached also (although they
have not yet been studied). Garstang has shown that
>11 t h · . resentlng
e evidence potnts to the conclusion that the
larval stage of Antedon represents tbe larval and not
the adult Stage of its stalked ancestors.
Ammonites 4J3O-'-- prOVl·de evl·d enee lor
r ...
devtation III
ontogeny. The schemes of phylogeny presented by
DEVIATION SI
Hy.tt and Wtlrtembcger present the following
situation. Microccras dcnsinodus has a spirally
wound shell of which the four inner whorls :arc
smooth, the nen two are ribbed, and the outer whorls
are knobbed. The inner whorls are of COUtIt: the
earliest to be formed, and 10 the anim.a.I Ms pilled.
successively through ontogenetic stages in which the
shell which it made was first smooth, then rib~
and lastly knobbed. So on tbe theory of recapitula-
tion, to quote Lang, 'it is supposed that e anceunl
form had a plain sheU, and that, during lUi eYOlution,
the stock or lineage from which M. densinodus arose
acquired, first a ribbed, and then • tubercuhu:
ornament'. This example is typical of >11 the argu-
ments which recapitulationists seek to base on c
study of Ammonites. Kow, why should the ~bbcd­
shelled stage in the ontogeny of M. dens'nodus
represent an ancestral adlt/J stage:? TJut an~
may have had a ribbed-shelled ear!] ontogene:lc
stage, but there is no evidence at all as to ~t u.s
adult state may have been like. Indeed, In JO~
. f Ps·,I~~' Spath has shown thu the nbs
SpeCies 0 ~ -, . ' CT
make their first appearance: in phylogeny 1.0 the ~;
......
u::
whorls, i. e. in the euly suges .of ontogheenYnd"
. furth er eV1·den co of thl> .~I r- co . I .
P • . < assumpbOn of a pr<V1OUS Y
'There 15 no necessity lor . .
. berculate Ammonltes., or
existing costate stage lfi tu before a stout-
of a slender-whorled evolute stage make them 6t
whorled involute stage, except to
 DEVIATION
Pb'tflll1 cycles and lineages; for the evidence gener_
at I . to some indifferent root J common to the
,1Iy pc"n" . I d
mes' These arbitrary eye es an lineages
t\ro extrc· f' I .
" on the theory 0 recapltu atlon the truth
are bascu
of which is taken for granted; they do not prove it.
I this cannexion, it may be recalled that these
li:eages may involve a reversal ofthe geological order,
which is of course absurd.
A similar conclusion is arrived at by Franz, who
points out that the smooth-shelled snail Paludina
neuma)Ti probably gave rise in the Pliocene period
to Tulotoma, which has ribs on the outer whorl of
its shell. The inner whorls of Tulotoma's shell afC
smooth, as arc the inner whorls of the shell of Palu-
dina) but none of the characters of the outer (adult)
whorls of the shell of Paludina have appeared on the
inner whorl of Tulotoma's shell, although they have
had all the time since the Pliocene period to do so.
Am~~ni{~s and Paludina therefore present evidence
of slmilanty between young stages and divergence
between adults.
\Ve may now consider a case which does not pos-
sess the evidential value of those discussed above
because it is based on a hypothesis instead of 0':
pr~f, :ut it is a hypothesis which no facts contradict
an w lc.h does fit all the facts consistently. It con-
cerns the origin f th . of the origin of the torsion of Gastropods h~s two
th ho
roug 180 wh' h .
0
e torSion of the body and shell
(ropod M II IC IS SO characteristic of the Gas-
o usca. Such a form as the lim et dev:.;e:;;l;;o,.p.;.s
DEVIATION
. 5)
IOto a more or less symmetri 1 VeI'tger Iarva which .
su d denly undergoes a twist th h 8.
f . roug I 0) the pro-
cess 0 torSIOn occupying two or three minutes. Now
Garst~ng has shown that there is a great advantage
accrumg to the larva from being twl'sted'
, lor I w hereas
befl')r~ the torsion it was not able to withdraw ita
head IOto the safety of the mantle-<hamber, it can
FIG. 3. Stages in the denJopment of the limpet, Ibowing that the
animal starts with being symmetrical. and lubseqUl:'nt.ly uncktgoes tonion
while still young, resulting in the twisted structIJre which is chal'2.cteristic
of Gastropods, After tonion the larva is able to ..-ithdnw its head into
the protection of the mantk-<:hambu. (From GMSung.)
do this after the torsion. Torsion thus appears to
be a larval adaptation, and Garstang is of the opinion
that Gastropods have twisted bodies because twisting
appeared as an evolutionary novelty in the larva of
their ancestor. This twisting would therefore be a
caenogenetic character which differs from. those
which we considered in the 1:lst chapter only 10 that
its effects persist into the adult stage instead of only
affecting the larval stages of ontogeny. This theory
advantages. In the first place it does away w,~h the
difficulties which arise in any attempt to explam the
_
 DEVIATION
54
.' of torsion on the assumption that it arose
nglO
°d unng the adult life of some early
.
Mollusc, No
adaptive significance can be conceIved for such an
event, whereas, as we have seen, the process does
appear to be adaptive if it takes place in the larva.
In the second place Garstang's theory provides an
explanation for the total lack of forms showing any
intermediate stages in this torsion. Torsion can take
place easily in the young larva and twist the body
of the animal through 180° at one stroke, but if it
took place in the adult it would have had to take place
gradually, providing intermediate stages. That inter-
mediate stages are not a priqrj impossible is proved
by the fact that intermediate stages in detorsion in
I~t~r Gastropods are not wanting. There is no possi-
bility whatsoever of mistaking these detorting forms
for the 'torting' forms for which we search in vain.
Lastly, we may consider some cases .in which
animals differ from others in the number of certain
parts. Nymphon is a Pycnogonid with four thoracic
seg~ents and pairs of legs. Pentanymphon is similar
but It has five t h · d .
'd'
It IS Iflieult to oraClc appc" ages and paIrs of legs,
. f '
conceive f
0 the extra segment and
r ur
0 legs ofPentanymphon having arisen gradually,
lor segments .h
P are elt er present or absent. or can
entanymphon be
N h regarded as the type from which
rmp on arose b ad
fOur see y gr ual loss of a segment, for
to
num\..-- m be the regular and five the exceptional
UCT 0 f segm .
ents In the thorax of Pycnogonids.
DEVIATION
Th ere remains'h t e »O5!1'b'l' Sl
I tty that the ntra legmtn.t
of Pentanymphon was added during the early 12 n
of on~~eny ,:"hen the material for the futlJre thorax
wa! diVided IOto five instead of into (our. St..ch a
larval variation would of COUrse affect the adult al
Similar conch15ions can be drawn (rom ot~~
examples, I have myselfobserved that in the develop-
ment of the chick it is pos!ible for the number of
segments, rib5, and vertebral bodies, to be greaur
by one whole unit on one side of the: body at COf1l..
pared with the other which was normal. I ..u also
plain that the discrepancy could not be ucounted fo<
by fusion of two segmen on the side with the smaller
number, nor by secondary splitting on the othtT lide.
It must be concluded that the proces' ofscgm<ntation
has split up the material of the chick', body into a
different number of segment! on each silk. In
Bateson's words, there has been a fre5h 'distribution
of differentiation' on one side. lfboth sides had betn
partitioned as was the side with one segment too
many, the animal would h1ve been companblc to
Pentanymphon in this respect, and for~ as a
result of 1n embryonic v.triation. It is possible th1t
the extra gill-slits which arc present in Hept1nch~
HexanchU!, and Pliotrema, over , and above the
number which is typical in Selachtan~ may be ex-
plained in this way as Versluys suggcsu, but there
are difficulties in this case.
Goodrich has shown that the position of the 6ns
 DEVIATJON
6
1 b ell transposed up :'Iud down the body 57
(fish h,ve e "h"C"
a J but that there JS no S IItll1g during
'" phy ogeny, .' .. Vlll
r The fin ames In :1 new POSition from
onlog en ). near:tnCC in ontogeny, Wh"IeJ1 sows h
• r.. that NEOTENY
ItS IITst :tpr-' • .., .
there hll:i~ been nO recflpltu!:ltlon; IIlstend,
•
"" hos resulted in the substitution of a new
•
progressive
O UR object now is to consider those cases in
which the adult form of an animal bears features
deVIJItion ..
posicion (or thlll ~(the :lnccs~or. ewertzow. has pre-
stilted similar eVidence, which he uses to Illustrate
by which it resembles the young form of its ancestors
"
or to put It the other way, those cases in which the
.
young features of the ancestor have been retained in
his principle of archnlln.xis" .
the adult stage of the descendant. Interpreting these
The evidence presented In lhls chapter proves that
cases in terms of heterochrony, they imply a rtlati'f.,t
\'on Bate's principles as expressed in his 3rd and
retardation in the rate of development of the body
,.th I:tws h:n-c a wide application. The origin in the (soma) as compared with the reproductive glands
c3d)' ontogenetic stages f vari:ttions which are (germen), SO that the body does not run through so
ontogc:netiC1l1), perm:U1cnt (i. e. nA-"c t the adult also) many stages in development in the ontogeny of the
is pro\·ed. These v:ui:nions difi'cr from caenogenetic descendant as it did in that of the ancestor. This
\':J.ri:J.tions merd)' in that the buer onlr concern the state of affairs mar of course be brought about in
elfl)' lages of ontogenr. But this difFerence amounts man)' wars. \\fith the nlte of development of the
(Q nothing but the lengths of time during which the soma remaining constant, this effect will be produced
inltrnal factors ontrolling the variations :lre acting, if the germen is accelerated. Here belong the wdl-
!IS would result from alternti ns in the intensity and known cas~s of p.udogtlltsisJ of which the most strik-
~rl~ilY of the tietors. NC:lrly all the 3ses mentioned ing is that of Polrstomum integerrimumJ which has
In thiS h:J.ptcr h3ve been:\ duccd to prove the theory alreadr been mentioned, The same phenomenon an
" I:ttl
f r tC:lpltu " Il, und It " hns been my t:1sk to show be obsen'ed in ~liastorJ one of the flies, Here, the
thllt the:' do II t prove it. \Vhat they do pro\'e is larva or grub produces the next generation ,,;thout
embr)'O '. ' 'I " " e\'er developing anr further,
me Simi 3nt)' nnd repetition of chal.lcters In
I n other cnses) the acceler:nion of de\-dopment
(CrTUpoIIJi"v .,\ st:\,ges f th "
e Ont gCllIes f :lIlcestor n.n d
I of the germen re.lati\-e1y to the soma is not ~ great,
ll"SCtndnnt " ) wh'th I reve,1 1s t IlC nffil\lty
"" between I"ff'cr-
('nt nnmmls b . I" h " 1 I"k
and we get nn :u\lm:t ' bl,...:.torn:t
I·C ."m .- whtch can
'd I 'I Ut supp ICS no evidence us to whut t e . . . . lan'31 -<;,t:J,rrt known ;IS
,I II ( nnCt:ilrttl C fl\\ \\'";15 like.
,...
become nmture wh('J} It IS 10 Its 0'
 58 I I lich a -case is usually called neo/ellOliJ
the axe ot . . I '
L' all e is of p:trtlcu ar Interest. The axolotl
and tulS . .. . .
, 1'ts external gtlls and gIll-silts, and thIS i,
rer-llns
number of other newts-the so-called
Lat a
c:<:tC[ IY Wit.
',branchiates-have done. But whereas the
P crenn • than those deSCribed above I for ,'f t n..
axolotl is only facultatively neotenous, since it Can
metamorphose and assume the adult characters of
Ambl)'stoma under either natuf:11 or experimental
conditions) the Perennibranchiates (including Typh-
Iomolge, Necturus, Proteus, &c.) :lre permanently
neotenOllS. There are other newts in which neoteny
does not occur, and it is clear that the ncotenous
condition of the axolotl is an evolutionary novelty
resulting from heterochrony. The Same conclusion
may be applied to the Perennibranehiates which must
have been derived from newts which only became
mature after metamorphosis, The phylogeny of the
Perennibranchiates, therefore, has involved the reten·
tion of characters (external gills, gill-slits) in the
adult stage which belonged to the larval stage of the
ancestor.
The Appendicularians present another case which
must be ascribed to neoteny. These little animals
nev~r get beyond the tailed condition that the sea·
squirts pass through; the A ppendicularians become
mature in this st atc, wh creas the sea-sqUIrts
'd an
others deveIop f urtncr
L
and undergo an extenSive '
metamorphosis.
It is easy to sec that a Condition of neoteny will be
NEOTENY
, L' d'fL
aCnleve I tncre 'IS no acceleration of ..'IL 19
d lief germen
or soma, but a rdatlvc:1y greater ret. rd.atio n in tht'
rate of development of the soma. uth C'a' .\\ c-
. . :tN '" I l't'
more lIltercstlng from
. the point f ,-iew of Ph,. I(l~tn
L .. Cl" • •
~(':ml('n I
:l.cce1eratcd the structure of the bod)- will be I('~ fully
developed when lhe animal is sexuallr maWf thll;l
was the body of the ancestor. n lhe other halld, if
there is nO acceleration of the ratt' f dc\'cl pmt'llt
f the germcn, but nly a sl wer f,Ue of SU(Tc~sion
of ontogeneti stages of the soma) lhe bod) "ill he-
llO less 'well made' for retaining the lan'31 piJn. '1 he
principle here described and to ht' illustrated ht'1 w
is that to which Bolk has applitd the terrnjlJ/t"Ii••I'
1;011. His ideas were based on a study f human
evolution J but as they may al~o he applit'd to Olhrr
animals which do not possess:t 'rO<'tal' staAt"t it \('('111
advisable to rct:tin the marc <: mprchtll in' tefm
neO/lUl)J which fi fms the litle of this !l:lpt('~. C:lr·
stang's term p(l(fdomorpllosis Ill:!.)' he npplted ~o ~Ildlldc
phylogenetic progress by lle t 11)' rllld dCVI;llIOIl.
Balk h:ts shown rhat mall)' of the fcalllTrs of lhe
:tdult structur of l11:tn show rcc;crnhbnces ~o th Sl'
of the embryonic structure f Ihe anlhro.I)()ld ap~ ..
These features include Ihe relalively hIgh beam·
weight, the position of the ~ ralll n Ill~gnum and Ihr
cranial Aexure) the retarded losure f th~. sUlUr,C'
between lhe bones f the kull, lhe d~nllll II J IlC,
(rtnog'''lhy), the h"rlessll'", I
flatness 0 r 11C r
I J~ce
 NEOTENY
h rght colour of the skin, and a number
60 bod)', tel NEOTE.NY
Ine 6.
( other features. . . it is not formed and the face is flat, the two eyes can
o The aXIS "(,he head forms a nght angle with that come to the front of the face and let their fields of
0
in the embryo of all mammals (and of vision overlap. This enables both eyes to be focused
o( the "un k )" on one object, and Elliot Smith has shown that this
I II ve, rebrates1 for that matter) and thiS
ncarra •
bend
"kn own a5 the cranial flexure. ''''hereas . In mam-
IS
I thef than man the axis of the head IS rotated
masO h " I'
during later deve/apmen.t s~ that t ~ anl1~a shead
points in a direction which IS a continuation of the
line of its backbone, in man the cranial flexure is
retained so that his head points in a directioll at right
angles to the axis of his body. Since the direction in
which his head points, i.e. his Jine of sight, is hori-
zontal, the position of the body will be vertical; and
so man's erectauitude is associated with the retention
during ontogeny of a condition which in other mam-
mals is embryonic and temporary) as it presumably
was in man's ancestors. The crect posture of man
is, in Balk's view) a consequence of neoteny of the
shape of the head.
The retention of the cranial flexure accounts for b
the position of the foramen magnum, the hole in " b' L . 1ngk which die head makes

t~e. skull. t.hrough which the spinal cord enters, Fie ~ A leries of xtlIons
. "T.. I owmg t,.,
b
L·
b ro buman "",mg; (,'
with the trunk In a,embryo dog; ,e~ r}
dult d",,"
. pl't'frvN in adult
,
d, adult human being. The embr)"oDlc curntu~ "
",hlch POSitIon resembles that which is found in the man. (From Bolk.)
embryos of the apes. The retention of the embryonic
condition also aCCOunts for the flatness of the human development of the power 0 f accurate.
vision
(
has
. h evolution 0 man.
::lce, as compared with the elongated muzzle which played an important part III t e f h human skull
IS found in othe I ' h" "( "
r mamma s. r IS feature IS 0 IIn- The sutures between the bones 0 t ~. In
porrance for a . I f arly thirty )ears.
do not dose until the age one.
• mUlZ e separates the eyes, and, when
•
 62 d ther m:tmrnal. these sut\.lres close much
apet an ,0 birth and when that has happened 'he
toOner IJ ler '. ,•
t increuc in "I.e any more. J he human
skull c~nno ., . r
thcr hand c:an ,"crta e 10 81ze lOr a very
th
.kull on e o . bl.
. all',er birth , 2nd thl5 ella es It to provide
long time
tion for the large volume f the human
;lccomrnoda • .
· Another interesting feature IS• the absence of
bnun.
large brow.ridges ( uch as characterIze ,he skulls of
d I, anM and of eander'hal man)
'u """
"' adult man and
in foetal apes. These brow-n.d.. ¥cs arc subsequent
deveJopmen in the apes, and I homson has shown
that they ful~1 a mechanical function in buttressing
the upper jaw against a heavy lower jaw. Man is
neotenou, in not having developed them, the but-
tressing function having been taken over by the
venical wall of the forehead, and the necessity for
thi, function having been reduced by the diminished
site of the lower jaw. It is because it has no brow~
ridge, lhat the fossil Piltdowll man is regarded as
close to the line of modern man's descent, and
Elliot mith has pointed out that the occipital region
of th~ Piltdown skull bears strong resemblances to
'hat of a young ape.
. Apes when new-born have white skins in which
pigment only becomes deposited during later develop"
ment, and the same is true of the egro. In this
respeet the white races arc neotenous for they retain
'he emb . ryome . COn d'IliOn
. of other forms.' One of the
mOSt Interesting cases of this kind is that of the hair,
fOHNY
6,
for Bolk has shown that a progre. ive te:riel in reduc.
tion can b made out in the monkeys, :1.pe1, and man.
I. The monkey is born with a tomplete COvtring
of hair.
'2. The gibbon i. born with the ht.ad and back
covered with hair, and the other re~ionJ are
covered later.
3. The gorilla i born with the head covered with
hair, and the other re ,ion are partially
covered later.
4. Man is born with the head covered with hair,
and the other regions are 'CUtely covered
at all later.
(I t is to be noted that the lanugo which form. a
very fine covering to the unborn infant before being
lost is also present in the unborn apes. Further, the
lanugo is retarded in man, for he has not completely
shed it by the lime of birth.)
'I'h is series shows that the neoteny of ma.n a!
regards hair is associated wilh a progr~live relar~a­
tion in the rate of its development. ThIs retardatlo~
in the rate of development of the body or soma, It
will be remembered, is all that i! required to produce
the other human features mentioned above. It there~
fore becomes interesting to inquire whether the rate
of human somatic development is reall~ slow ~s
d with that of other mammals. fhat thiS
compare bl hich wa5
actually is the case is proved by a ta e WI .
.
given . Chapter III • Bolk has been ab e to give
111.
 N.cUJeNY

64 .. aIproofof this by a study of the developn"

addJtlon 'Ik th ht ~EOTE~Y 6-
th
ofte< . In the apes, the mt -tee afe Cut direct!
I . c y
,
This case has been described fairly fully to serve
hlf. th the 1St mo ar IS cut SOOn azter the 2 d
al[a , fh. h as an example of the phylogenctic effect which
remo Iar, a nd the replacement 0 t e milk-teeth
. the n
P heterochrony can produce br allowing characters
takes place accompanied by the cu.ttlng of the 2nd which had been embryonic in the ancestor to ~come
and 3ed molars. In man, the cuttIng. of the rnilk_ adult in the descendant. Two more cases will now
teeth is only finished two y~ars after blr~) and this be considered, from other kinds of animals.
. fl Hawed by a pause untIl at five or SIX the 1st
:01: is cut. After this, the milk-teeth are replaced, c.hated band

and not until this is done does the 2nd molar appear.
The 3rd molar may be cut after the 2nd, but its
development is often so retarded that it is not cut
at all. Indeed, retardation characterizes the develop-
ment of the human dentition as a whole. Associated
with this retardation as Balk has shown is the
prominence of the human chin. It is interesting to
notc that Neanderthal man had the simian mode of a b ,
tooth-development and lacked a prominent chin. , be· he lan;l of;ln Echln~rnl (<lIj ;lnd
FIG, 5, A eomrarJJo~ . ~\\~n I h ;lnd () The bi.nd. of cilia.
It may then be safely concluded that the rate of the form of;l typical prlmlU~ C~:1~le.Junl fold". of the latter. ~ do
development of the human Soma has been retarded. of the former correspond e.:u Y._ d h aide \W:II" (, duna.1 V!CII'.
the adonl Nnd and the e.ndofitpt:" a a n . •
On the other hand, the reproductive glands have (After Gantang.)

probably not varied their rate of development, for I k for the trace of the
Garstang was the fi rst to 00 , d f dult
the human ovary reaches full size at the age of about f h t brates in early mstea 0 a
ancestors 0 t ever e h ~ sed his attention
nve, and this is about the time ofsexual maturity of the stages of invertebrates i 3 nd e OCt"rfish sea-urchins,
fEh'oerms(sa
d I
apes,and presumably ofman's ancestors. Thehuman on the larvae 0 c In h d that if the ciliated
b &c) He s owe
bodyisJ ,however, not ready for the reproductive glands sca-cucum ers, .. , .) r a sea-eucumber
tofunct ·1 . AUricularia 0
Ian Untl several years later. The retardation IS bands on the 1arva ( 1. groove between
due to the t" f . . idges eavlng a h
tant pan inacr IanI·0 hormones which play an Impor-
,
were to rise up as r r
. these rI.dges were to u ,
se converting t e
egu atlng the speed of development.
them , and If
3666
,
 " ....... '~ . ..
66 o ,tube a structurc would be produced
oo'Ve lOtO, I
g' o h ali we relations of ,the vertcbrate nervous
whlCh as,
. cluding such detads as the neurenteric
sptem, IO dO fio
I Not only this, but the two mo l cattons of the
=' bO
verte ra
te ncr'VOUS system which are found in Am-
phioxus and ali higher forms on the one ha~d and in
sea-squirrs on tbc other, ca~ be ~~sed on dlfferences
which are found in the dlSposltlon of thc ciliated
bands on diffatnt kinds of Echinoderm larvae. This
theorr of origin of me vertebrate nervous system has
se~-eral advantages. In the first piace it avoids the
difficulties which bestt any attempt to derive it from
tbe existing nervous systems of any other inverte-
brate. It also agrees with the principle of neurobio-
wlS according to which a concentration of nervous
tissue takes pIace in the region of greatest stimulation.
lf tbe ancestors of the vertebrates had crawled about
on tbeir ventral surfaces like most invertebrates, one
wouJd expect their nervous system to be ventral, as
that of most invertebrates is. But the nervous systern
o.f vertebrates is dorsal, and it is precisely the dorsal
SI?e of the body which would receive tbe greatest
s~mulatioo in a form swimming freely in the sea thc
stlmuli being the rays of light penetrating thr~ugh
fro.m t~e surface. t ot ooly is the Auricularia free-
SW1mming, bUI IOr bears an unmlstakable resemb Iance
o
toI tbe Torna ' Iarva o f Balanoglossus, and Balano-
' fla
gossus ,sanudbd o of AmphlOxus
o d
Ih e early ve I b n ou te relative an
re rates (or chordates as they may
NEOTENY
'7
more correcdy
, be. termcd). But the rcc"n'bl .... • ance
betwcen t.he Echtnoderm larva and the cbordate goes
furthcr stili, for the former has an adoral ciliated band
formed partly fr~m the i.n~er layer of the body, and
in a correspondtng posltlon the chordates have a
ciliated band cal.led the endostyle, which is looped
in the same pecullar manner as the olber. The middle
la)'cr of the body arises in three tiers or Stgmentl in
the Echinoderrn larva, in Balanoglossus, 2nd indica-
tions of this tripartite arrangement are prestnt in
Amphioxus. FurtherJ the body cavit}" of the Echino-
derm larva is in communication with the oUlside bl
a pore, as in Balanoglossus, Amphioxus, and Stvenl
other chordates. In fact, if the nervous system ami
endost),le are formed in the wa}" suggested, a\l lh:lt
is required to turn the Echinoderm larva into :I
chordate is the formation of the notochord and the
piercing of the gill-slits.
Now the Echinoderm larva undergots an exten-
sive metamorphosis during which the generallar\'al
form and symmetry are Jost, and becomes the adult
starfish sea-urchin, or sea-cucumber, as the cast m.t)'
, . ' l and pecuhar
be Their adult structure IS so speCla
o f ardO g the adult form
that nO one would dream o reg In • Il
....... 1 tO an)·thmg :lt a .
of any Echinoderm 15 ancesu ..
• ID th hordates were
Nor indeed is it proposed ~t ode c s the)' no\\'
derived from the larvae of Ecbm . ermi"hose aduhs
h been anlm3 s w
are. But there must ave . h the aduhs of
were less special and pecuhar t an
 68 NEOTENl"
existing Echinoderms, and whose larvae rCSC bl NEOTENY
" E . m cd
those of existing chlIloderms. If,•
then J by h Cero...
I and an abdominal region of about ten segments
chrony the larval form ofrhese animals persisted which usually do not bear
. any legs " Now th ear~ I
. and
they became sexua II y mature
•
In this state
J Sue h
of lulus hatches as a httle animal with a head which
ncotenous forms would .provide exactly the /lce CSsary seems to be composed of the same number of s~g.
.
ments as that of the insect, and a bod)' of rou hi
matenal for the evolution of the chordates 0
n t he
Jines suggested. Indeed, the general form and sym. a do~en segmen;s of which the first three each ta~
a pair of legs. fhe 4th and following segments of
the body are not devoid of legs but they are retarded
in development and ~o small that they do not pro-
trude far, as Metschlllkoff showed. This six-legged
larva of lulus eventually develops into an elongated
adult form composed of several segments and bearing
many legs. TOW, if the retardation in the develop-
ment of the legs behind the first three pairs were
increased) and the larval number of about a dozen
body-segments persisted into the adult stage, there
would be formed an animal like an insect with the
Flc. 6. Tilt, bmll form of a M 'm. . legs behind the first three pairs either reduced to
itJ 1=-1 teIt bb ) pod lOOn after h;ltchmg, showing
m nee to an In!l«1. (From \'om Rath.) vestiges or vanished altogether. It is therefore of
interest to find that there actually are insects in which
rnctry of the Tornar' d .
develops' h la 0 persist when this larva vestigial legs are present on the abdominal segments
La 1 Into t e adult Balalloglossus. (Campodea, lapyx, Machilis). These insects have
st }', we may turn t th . .
be drawn b I 0 e companson whIch mal' no wings) and on the other hand have preserved
e ween the d I {; features which the other insects have lost (such as the
the larval {;o f a. u t Ofm of the insects and
rm 0 certa M . . segmental arrangement of the reproductive gl~nds),
(the millipede) Th In ynapods such as lulus
sists of a he d' b' e ~dy of the adult insect con- and they may therefore be regarded as an IIlter-
aWlchlsmad f' mediate stage in the process of helcrochron)', :llId of
segm~nts SOme f h' e up 0 SIX or seven
) oWlchb' the evolution of the insects from ncotellOUS brvae
of thrce segmcllts each ea~ Jaws; a thoracic region
of which bears a pair of legs; like those of rhe Myriapoda.
,
•
 , ~tl ;;1'1 ~r~l· 'l'l Rl-~
• • • I UK 'I'll RKOUCl'lQX •
~~ ~t :1, Iii 'tU('~troh \\~II
,( the (',nt ' _r-,t.g~'S f den~lQrll\cl\t of th,'~c dnlllU! . l
, h' "''t'''-~!l'''n.,• 1 . lo,' ••

, .
\ ,
~1\
•
I.
fhc. I '~CSS'OI\. th o..t these. \'"CStilYe$
'e
is 'h t'.ft'\Nl;:
...._ no.
~.t\: ~I, th(' u • \'"knee ot• c 1'rt":s'Sltlg b.\ck of aJult , n lrt I
-1\.'"'
, , '
tt"N! lnttl the ung smgcs of the d...,,,,, )
.~l ... nu~t.
t,""t>' btl'"....,· th< 1 he S.Ul\C IS true f the tooth-band lnd 5011\111 h
>tot.., ,~~ltt it. '} <oh- . . d' (rtt
wbu.:::h "rt":. \In ~n the cmbr) of the \\ halC'-bo.nc
i < \' h:tlcs... hcsc "nlnuls lire toothl tn the adult 1
. ' 1 lnu
thcir cmbr me tttth ~I'~nt the cmbr) nic nJ'"
rion ( tha m:a..mm"ts. th.tr n"'1.mpl ~ the N.
of the "'Ou"",-, "ttl while its adult h onh' h\
the IN I'~~isting in the dull s~ . othtt
"rds·therudimcnt . . fJJ. n'sQr1!'1.ninanbn ic
~.

nan ",1\ •• It In the llduh s... ;-bile it is ~nt

in othtt :lJult mamnuls" I.nJ the ruJint<:nt . . the
h.. idn"y or pron<phros "'hicll is roN in til<
~mhry '·sa.... of211chordJ~~ in(h('sJult
i n < fi'h but is 10<, in lb< ,dult . ,11 « t.Orm,s.
"·e ~) then) th~u :lS in th~ a~ . d~-ia' tht:
insQn(xs in which stnJdUttS ~ nJ.uttJ to \ ~ .
ob<y''On Bau's b of th< £T""~ dq;'« ""'In-
bb""" thot <Xi, b<twttn. OJ 'ng<' • Jitf«<nl
uimals than bct.ccn ) utJ • ult suges or
between aJult Sbg'd iAu:r Y.
X
AD LT HRLHIOX
'DER this hndi ' f t indud< ... ,t ~ 1'<>\
•
•
•
U Ol\ly.
descr......, h_.'_
1~ WJU"OJI
.L_
UK;
• ~_ .J..onnm<non ...
sp<ciaI as< 01 me " " - ' _ . .
'-'·lion· As has
to i" \,I'It:~-
....
b«tI

•
 ADULT VARIATION
,. ___I I--fore the reactlons
" hW"Ieh result in th
strt'SKU ~ I . e chrony, and produces only small phylogenetic effcl..t ,
llppearanec of characters are set gomg Some time
,-
before those characters appear: The adult form is
"
being pr
cpared during the earlier stages of deVelop-
men.t We cannot draw any hard. and . fast line be-
tween the characters which, substituting themselves
for others in phylogeny, appear early in ontogeny,
and those which appear late. However, the later a
,hmeter appears in ontogeny, the smaller as a rule is
the change which it produces by its presence. On the
other hand, a character which appears early in onto-
geny has time to produce more important changes.
So while under the heading of deviation we include
variations which have produced fairly large phylo-
genetic effects (such as the divergence in evolution
between different families, orders, and classes of
animals), adult variation deals with the differences
between individuals, varieties, races, and, probably,
species. Darwin himself drew attention to the fact
that 'slight variations generally appear at a not very
early period of life',
To this chapter, then, belong the slight differ-
~nces ~tween animals, and genetic research tends
'0
increasingly Show t hat these dIfferences
" II Th e bo nes are vestlgtal
are con-
"oiled by M en d eI"Ian factors or genes, and that they
may concern any and every feature of an animal. No
~seful purpose would be served by going more fully
Into them
.' Th e su b" " of one character for
Slitutlon
anot her In the aut
dId oes not usually involve hetero-
ADULT \'.\RIATIO~ 7S
Just as, however, cacnogenetil.:. variations may be-
come phylogenetically important by undergoing
heterochrony, attention must be paid to the possible
results of heterochrony if applied to adult variation
and this will form the subject of the following
•
chapters.
XI
VESTIGIAL STRUCTURES DUE TO
RETARDATION
H EN as in neoteny characters which had been
I
W larval or embryonic in the ancestor become
adult in the descendant, the original adult characters
of the ancestor tend as it were to be pushed off from
I the end of ontogeny. They arise tOO late to be fully
formed by the time maturity is reached in th~e
animals which have a definite adult form, and In
consequence such characters become reduc~d an.d
vestigial. Retardation of structures to vestiges hiS
therefore the other side of the picture presented y
" I"s vestigial on the
the phenomenon 0 f neoteny. !-I aIr
body of man, as are the molars, especially the ,I~st
which are often not cut at a,
"II bone
in the skull of the axolotl, and the maxI ary ,
h'ch undergo meta·
which is present in the newts W I . .
"I d It form IS absent In
mo'phosis to the terrestna a u ,
" "h permanently neo-
the PerennibrallchlatcS whlC are
tenous.
 "rSTIGL'L TRUCTI,;R£
------L.IIII
If is possible to interpret these cases in (ttnls of HYP£.lB.IORPHOSIS ._
th< pb<n<""""'? of inte=ntili1J in the gypsy Uloth rqoroJueti\'t _ bods' dd~yt:d rdatiYdY to tIut .
bor noric-:! lht In 2; nornu.I fem2..l~ moth, ~ Il1ak- the- bodr-clunatts.. '1M cases b) be ~ httC'
~ . ~ gwcs 2l"e (00 slow [0 produ~ the appear. confonn to the principle of'o==p . " '.J
:anct of any male dwzacrs before the tune of matur- and of p.~00n as ~ .0;1 bor F~"t. •

itT mim: :and dettJopment crases. Chan ers rna Achthc:res is :I 0"UStIct:ln ~ _ e- CO('("-
~e ~ by the oV~--reurdat:ion of the ra(~ pod'S, which in i dC""dopmern ~ a 5(>-
of action of the f1crors wblch contto) them. The aJled 'cope-podid' sage, in which it oalv dilfttS n'"",
dft'CtS of such heterochronl ,,;11 not in themseh.cs a nornul adult copepod in th2 Ihe- scgoau . its
be' imporwtt in phflogeny, but the)" will be aSSOCi_ thoru 2fe not so clC'2TJ ' mnko:i, ud its limbs U'C'
[e-wer Uld mt:tlkr. The adult AchlhC'n:S 'as the ~ull
:ued with neatenr, ~d this, as we h~ve Sttn, mar
of further de-vdoptnC'nl is puniti<: 2nd ~~.
and often does luve Important results In C''Ulution.
Another comp:ulson of the sm'te sort ._ hi N noJr
XII herwC'en the '2 uh Mysis (the 'opossum' shrimp
which hu for- ed J~ sod thC' so-alkd 'm... ~
HYPER MORPHO I
in the c\"dopment of tbC' lobsttt. The adult lobsttt
TF by he erochron~~ characters which appor in the h2S lost one of the pron_ och' the- ._.
~uJt . ancestor are ret:trded. so that the)· do not fork<d l<gs.
appear In the ~escendllllt br the time deVelopment werno\\"s principle of 1fts.bol}' is a moJinanon
cea~ they WlJl become \'estigial, as shown in the ofth:lt of'o\"USteppin_ '. In the on "" ~.:. ,,-nttov'
preYlous chapter. But if the time whe.n development distinguishes 2D e:ulier period of morphogt'n~s
~tops is rt/tl/;wIJ delayed, it will be possible for the during which the \"".lriou'S strununJ chu:l\:ters ~ th~.
esunda.nt to add characters on to the adult ancestnl devdoping :lnim:u :lrt moulded, and :I b.t('f l~rioJ .
stage. Referring again to the example of inter- growth in which the :lnim:ll incros<'S In. SIU unlll
• it retches the :tdult st:uc \\;thoU[ und~ng further
sexuality in moths, male characters do not normally
a.ppear by the time development ceases but if the modi6clt:ion. He rqr.trds it lIS passiblC' tOr ~ d5.."\"D-
.
tunc of marunh.· . ,
J IS postponed, these male charact~
• d:lIlt to :tdd :I. ncw sage on to the wt s~~ of ~~­
phogene:sis of the :l.n('('Stot. :\s 1ft ilIustn~ thiS
Id
~ and do make their appearance, This addition:ll
e\dopment
.
h
,or lJPumorphosjs, mal then be ex-
pttt<d In "'ses Were
h h
t e r.lte of devdopment of the
I
principle we 013" me the devdopmenl the ~~
snoute-d fish lkl~ne-. In the "dult, Btlone b"J.s

•
 /fYI" ,'~f()lIf',,(al"
7' ,
,I,,. lII' ," a,,,J Ihr I,;w r ).iW Vtry, "HIt
• •II d'JlH'<tI.,1
') u, WI' ,. Hi. \f 'J'" f' II (J J
1'1
.raj( Hi mIJ'I1"O~r" ••, II. J w~ ar IJt,Ih
, \.I.1"h .
I'll in early
' Urnt'\I'. nil Ih" "UC::",ilt~ n u f bypnllKlr..
. . ~, "" Ii
•hl}rl, If i" ht' Gnal 'tAU ()( mfJrph I JJ,(C'l flillJ f} 'Ut.-h I' hmit may .ht" ,aid v, "m(fAm II) J f....
- ~LI. ..I_
( U 1""llKl t_
• n.h at J'itl)((JCfU', At a J~, -, .t.. U~, the I'Jwcr jaw h
o I tt{.lfmulatlfm
. I)f adUh(fJr fin.al rtv b
J1p t.'lt(r~1 )
o( IJdon. d""!I.1 , Kfntly. p,odu.",U Ih '1'" ·.r. 'la",et In lh (Jnr"l{tny of th.. dnundatlt. h it: It, I(
whi,"
.nre iI" K'tH 4! "r tit final ltaU (j Il1f.1rJ'ht.... "'lIed, ~hen, fhat the 'rtupitul",'vJf1' i. hilt i t If, name
~c",i," .n',lher r..h-I f rniromphuJ, Laatly. th woulo,mply an ahbrevial.,.j (i,t, 1(.«1...."'.4) r
ion, hu a .implc ref ilion_
l-

upi ,j.w', BcI'J~, .1..., , I',nu. te • to I"OOU" Ih

J(lt'll n{Jul.·d (JJNf..hlJlm or ," finOiI .riilH or mfJrrhu-.
"'''W''', whie h then t1Jaru. hy .,rowth withrJU
(u,lh" "",.),r.,.ti,," to (o,nl Ih dult, f will
be n'jli,c4 th., , w 'WJw', an'boly dilli ,. (rom
'bV , 1'I'I"nU' ,,"ly in Ihat it i, Ih r.Ilal ataU' o(
llIo'pho en "ill t<ad (j th 0 fi"itiv adulla"B<o(
th' .n' W whkh ia "" cd lh'oull)' ill th ontog.ny
ofth dc" no.",. /lUI i" ilhcr," c,lh. LaB' which
Ih. OraW'O'"t In a Ih,ou h i, Ih fin.1 '''g' o(
or
'dtv ItJpm""t' (he: ant lOf, ror d v l0l'm nt, i.e.
.'fUL(Ufifl altcriirifJn, dOt. no uk I,I:H•.C during
!i<:w r(lOW', Il<,ioo o( growlh,
A ,II.v 111,,1)0 h "ral1< '"0 Sew rwJW loy lire"
'" Ih ("',llh' the aouil ""II or final .taB of 1110r-
~h'j~el1 <It of Ih .".."'" i. not p,tI eO b"k il1lO
,,,I,,r taM' of 0 v lopm III in Ih ' 0., <l1dal1l, but
h. "nIrJ~ ,"y or morpholl n .i, of Lh laller i, pro-
""B'o, ) h 'OUII "o"Oition 0' filial llOB' of mo'-
"bJKCnc.i, of th an"Ctilor i~ l,rClient unreduced in
h. d v 101"ncl1' of th d "cnO.nl bUI it i, no
Now j will be ol,vi'JUJ IMt the rrin( iplc o( hy
mfJrrhlj4)i. re"f entirely on the aMump'tr,n that a
•
nf'Jrm21 adult (rJpcpod wa ancalraJ to A(hthrra;
th:- a normal adult M yt.' wu anu: trOll IQthe t"r~
.I1J tho r.ah wh_ fin.lala~" of nmp!w>lrt....· lnd
ther (urc: wh,jt.C adult. were like thm.e of Lxoc.oct ,
and llcmiramphu. were anc«tofl of K(\t)M, I~ut
it there any evidenu: of thi.? It i. equally iblc
Iha "ornul <o""pod••nd Achlb...... had a common
"ncettor from whkh each h.ave dIVerged, and the
co".podid ..aBe of Achlhere. would be evUkncc of
.ffinily with 110rmal ",,,,,pod•. ,imillrly, My itand
the loblter may have had a wmmon anu lOr whit.h
Have ri1C to both, the mylilltage of the lobstCf how..
inKitlaffinity o My,i" Inthe~mtway,a,o~mQ"
ance"tor may have given rise to l:-.xocoetuJ, I ftmln",,"
phus, and Bclonc, the ontogenetic Itaget of Iklonc
b'lraying iu affinity to lhe.. r..h. .'
In the procell of these phylogenetic dlvcrgcn{(t,
A.hlhere. may b. regarded at having devfelopedb
1 oct.' tnc loblter urt er
furth r than norma ..orep n E. oetut, I

fJrlllflh' J than MY'li.i Ilcrniramphul further I an 'xOC

l!t C tcrmllllll HI:I~c of OlltoUCIlY or morpho"
 II Ypgl{l\'IOR P 1105 I S
80 . .
and Udone further than Ilemlrarnphu5. There is no
neccssity for regarding these forms as all on one and
the pme line of descent. I n other words) the in-
lanCes described in this chapter may just as well if
~ot better be regarded as illustrating the principle of
progressive devi~tion, dealt .wit~ in ~hapter VII. J f
lhis is SO, there IS then no Justlficallon for the view
that adult or final stages of morphogenesis of any
ancestors are included in the ontogeny of Achthcres ,
the lobster) or Belone. Instead , these cases would
serve as an illustration of the principle to which
J Eimer gave the name Epistasy, according to which
one of two related forms might undergo morc
modification in phylogeny than the other.
However, as recapitulation by hypcrmorphosis is
theoretically possible as the result of heterochrony
and prolongation of the period of development, these
cases arc mentioned for what they are worth, with
the (Om~lent that they cannot be regarded as proving
lIa~ckei s theory of recapitulation. It may also be
notlc.ed that the phylogenetic effect that hypermo r-
phOSIS may produce (if it really is responsible for the
cases above described) is not great.
XIII
ACCELERATION
W E have now to consider the last of the possible
results
. that heteroc h rony may produce. By
acceIeratlOIl of lh e rate 0 f action of lhe internal factors
ACCELERATION II
which control
. its formation • a charac'cr W h'IC h ap-
peared In the late ~eatures of ontogeny of an anCe'5tor
may appear early In the development of a desund2.nt
The embryonic or larval features of the descendan~
may t.h.en resemble the adult stage of the ancestor. The
Co~dltlo.ns required for llaeckel's theory of recapilU.
latlon Will then be fulfilled, but again with the rcnurk
thal accelerated repetition of a character in the antI>-
geny of the descendant does not constitute a pressing
back of a complete ancestral adult stage into an
earlier period of ontogeny. The fo\lowing examples
of acceleration may now be considered.
One of the most interesting features of the very
early stages of development of the chick is the pre·
cocious formation of the heart. Indeed, the heart
may almost be said to arise and to enter upon its
functions before the embryo itself is there. In the
development of lower forms such as the frog, for
instance, the general form and architectural plan of
the embryo is fully marked out by the time the heart
nppenrs. There can, then, be no doubt that in the
chick the formation of the heart has been accelerated
relatively to the rest of the body. The' ~eason' for
I this precocity in the formation of the hea~t IS not rar to
seek. The chick embryo is provided With an enorm·
\ ous quantity of yolk which cannot be ~ade use of
without a system of transportation, ~'hlch conve}
the yolk to the embf)·o. This srstem IS .the \"'aS~lar
,..
' h pump which nro\,dC\
system) and the heart IS t e
~
1';"
 ACCELERATION
8. . a I energy for t h·
e transportation. In the
the meC hanlc . .
h ther hand, the yolk, which IS present in
frog on ce 0 I d· ·d
' Iler quantity, is ene ose InSJ e the embryo
roue h sma
h earliest stages of development, and there IS
Iiromte f .
nO need II
ror a precocious system 0 transportation.
The chick provides anothc: example ~f accelera_
tion in the precocious formation of the Inner layer
of cells (endoderm, )'olk-epithelium) which precedes
the appearance of a structure known. as the bla~topo~e
(primitive streak) instead of resulting from It as In
the frog.
Another example of acceleration may be obtained
from a consideration of the tubes which grow out
from the middle larer of cells of an animal (meso-
derm) and place the space of the boo)'-cavit)'
(coelomic cavity) in communication with the exterior.
These tubes are called coelomoducts, and Goodrich
has emphasized that origin.lI), ther served to free the
reproductive cells which are formed in the bodr-
cavity. As the reproductive cells are not freed until
~hey are ripe, and they are not ripe until the animal
IS adult and has completed its development, the
coelomoducts must originally have been formed in
the last stages of ontogeny, as indeed they are in
many worms. I n other animals however and especi-
all . ,)
Y In the chordates, the coelomoducts make their
appearance in the early stages of development, long
before the rep rod · ce I
uctlve I·
s are Tlpe or even Iiormed .
Jn such ani I h . '
rna s, t en, the [oematlO of th coelomo--
ACCELERATION 81
ducts has been accelerated, and the precocit), of their
appearance has made it possible for them to take on
.
another function, viz. excretion or the getting rid
of waste products. This function of e:scretion mu t
be served from the earliest stages of development,
and in those forms in which the coelomoducts onl)'
appear late it is served by a different srstem of tubes
altogether (the nephridia) which arise from the outer
layer of cells. This latter system of tubes is not
formed in the animals in which the coelomoducts
arise in early stages of development.
According to Osborn, horns were present onl)· in
the late stages of development in the earl)' members •
of the titanotheres, whereas in the later eyoh'ed
members of this family the horns were present before
birth. This case has alreadr been alluded to in
Chapter IV where the principle of heterogonic
growth was mentioned. It was shown that the greater
size of the heterogonic organ, and consequentlf the
more rapid rate of action of the internal factors con-
trolling it) were associated with an incre:lse ~n the
size of the bod)'. It would seem, then, that In ~he
case of the titanotheres the accelerated nte of aetl~n
h h ogonic or~n to ansef
of the factors causes teeter
. . the case of the ere 0
earlier in ontogeny, Just as In th fer the
er the factor e ear I
h
Gammarus, t e strong . d d drawn
b i k Mehnert has In ee
eye becomes ac .. . tructures i. e.
th f t that progreSSiVe s ,
attention to e a c , d during phylogeny,
those which have become enlarge
 ,,(( I II IoIA'/I(JN
I, . M.nu RATIO'
"r IIf rolrl, r t"jU t 1)"1I'"cn 1 .Ind lu . '1
r
If II' I III II 'JI
-'Itrr tlWI
, '
they "lid . an,
(J
" a rt'\emlJlan"e tfJ Ihe adult as to the nrly "'.n','Ktnttl(
.
f JtVt I('f' (.. H1
. . . lItr:lllUflll't.
.HIKc, of the ante.tor,
"h",t' rulllpJ(\I .h"w lh:.t II I~JU"t RlI' """lIthl c f(J(
Many reupitulationi\tt would altO he di ~d tl)
,h.. "lI fr , ~hi"h III'P' rrd 1.lle 111 the unt(JHcllY fir ~"ribe Ihe re,emhlance between young deKendant
I !lIIH I(/f It} ullclrru o h tcro,hrol1y ~l1d 10 :ll'pt:lr and adult :tntcstor (whkh admittedly can occur, a
or
;~I~,y 1/. rltl' fJrltflW"Y the dtllt'tn<-!{Int:ll vit.:r vcr,,:., IIhown hy Ihe example. ~ivcn in this chaptcr) to the
In thr ,. (" r' th.. yfJUnU " .."('-' ()f Ih de Ctnd:ln, 'inheritance of acquired c.haruters·, We have alrudy
.hl,,,, ",r,drf .imil..,iIY with Ihe old tHlj.fC. rh;Hl with ~een thaI thi, mea", the conversion of an external
'lit younH III Hrl ,of III anu'.tor in rC~J1'U or lhe fac.lOr, whith originally caused a moditlcuion in a
dlJrolClrr 111 llur.,IOIl. flul Ih· flU '!crnuOIl of Ihell e tharaClcr of lhe b dy, into all interll:ll factor, which,
lll"'.u Irfull/CUIOl jll" i rYI he wnd u",iOIlI h:n I he cnrl y heing Iran mined by inheritance, produce. that me
tl"U e- Itl Ihe 011I(1U("1I1 0 lhe de "'/ld, III r(pre (II' modifi :uion in the ab~el\c:e of th2lt cxu~rn2l1 fa(lor.
1I1,llilijilluf '/tl' HJ,Ilplrlc ;",11111 rUH.CI;lor. We Ulnnot We may also repeat IhM s:uisfac:tory evidence as to
dllilll ttl o!JI,li" ill ornlatlon {Iii 10 th· Orlll or rhe the OUllrr 'I\{e of this phenomenon is still required,
,Ihilil/llt" IHllt_lor' (((JIlI:I 'fudy or Ihe YOU/1M' lIl.t(.;c' nnd we m:ty onc111de this chapter with a de cription
III Ihl' <.Jl'vclopmtllf rl:6 d· (CIlt!;lnl uult.·•• (tlclcr,l'" of en cs whith iIlu~trate the cart which is rcquirrd in
ullrd, (111(1 ntl W, ",Ive leen, il Illfly h' 111M
hfjll Ililtl f/l il,lcrprclillS'such vidence.
fI few dIM}j~lrr" her' rlml lhtl' ar'c :ll'..(:.c!cr:ll ti, hut In m:lll1m:l1s, tccth nrc formcd hellc;lth Ihe Rllms
Ih.11 i. Il, nnd Ihey may he cut hefore the)" ung anim.al is born,
II "illy I,t Il</illlrd (Jul Ih,ll tho who hclil'VC in SUlh Iccth ah w unworn c nieal Ul\PS or n~gr ,and
J IrH'l k(,j'. 'hrOt y II 1NllpilUbricJlI al'l' nnUIH(JlllCd 1(1 it is only oflcr birth that these cusps or nd~cs are
It,l' II, q'prlidllll of Illl' c'lfly (Jntll~l'IH.:ti 11,1~(,1I or
worn down (0 liar surf,llcs in the molars, as a res~lt
of the wear :llllI lear f gruH · l'llr I\llkcnlhal dls-
h t:!l1lflllhr Ihl' (,,"ly flllI(l~Cllrlil; 1ll,IHrll (lr Ihe
III lit'!'

!e'lrlllililll Uti If llin!' I ("peiltnl tlLIH'1i l'cprC!I('IlICd

covered Ihot in the arly unhorn cmbr)"o f t~e
l)uf:.!OIlK lht' Winding ltelh h:\\'e unworn rus,II'\s, ,IS
II udull 11l1l('IIWl", 'l"IH'y wOllld, Iht,'dolT, illdmlt
. , hter cmbf)'o, sll un-
I ,'hi. tlll4l'ltr Idilitc ClCIHIll'kll whidl w h,lv Il!ltd
Olle would expl'ct. II UI,11 ,I • f
I sho\\c{t flal sur accS,
tllu"!'ol'r IllflJ.tt(·."lv llrvitltiu/l, J I Ill,ly I)e rCf."llkJ hnrn, he foul1IJ thnl lhe Irct I r"
I '11 11)' flcfton.
11m.'
,
flq if Ih(')' hnl! b{'CII WOfll lO\\ , '\ rfSfnlS
hi! We' Wtll~ II, lillllll' \\111('. Itlilt. to "how Ihlll lht
H}'Olillfrt'I :lIlll11,1 p
I1ll'll, i.. 11 t,l'~C .11\ W I·11\ I1 t I'
lIy IIhl"rll 01 ~l drll ('lId\l1l1 did /lOI he,lf nil 1"'(',1'
 S6 ACCELERATION
(c::auue which it definitely charactcr~lic of the
:e-ecu of ute in Jduh aninul.; the embryo 'recapitu-
latet' the ground surfaces of
the teeth of its adult ancestor•.
An effect Ius been produced
, in tbe abtcnce of the C'Xtetnal
uUlCwhichnormally produces
i~ fOT KOunthal showed that
evcn if the embryo had the
habit of 'grinding its tu<h'
before it was born, they would
not weu one ana her down
fOT he teeth of the opposiu:
jaw. do not yet [Ouch. How,
b then, are these flat swf.aces
fK.~ 7- YIr:W"Jdltoow. produced? The teeth of the
oI*~ "no(' t '01Dugong are peculiar in that
"*0'1 ,.1ackJ<i • th~y a e
w' '0
~ +l _ . Jr.r) t1w
i
'"
-
r pcn...tra ed by cana" ••
tIiIf' - -d niIe:. the through which blood-ves5Cl~
_en' ,.,(t.n ... ..." d'
.. ..uas .. P' 'l(,. n pass,an Itappearsthat the flat
~-'la:aformc,dbr __· surfaces are due to removal
... 4o.a. (f~ JCljkc-IpJ)
. . of the lOOth-substance by liv.
Jng cells, which were brought and are nourished by
these blood...vascl,. I is obviow that the internal
{.actor which controls the formation of the Rat sur.
facet in the unborn embryo i.s totally different from
the ~nal factor (friction) which produces the same
~esult In thc adult. Can anyone suppose that friction
In the adult has had anything to do with the origin
ACC£LERATIOS
>.
of the internal factor which COIUJ'oh , _
of the toOth in the unborn embryo? To ClClrpI:
a concluoion would be '" faD inlo , : :..-
'post hoc ergo propter hoc' type of ~
It is interesting to compare • ~. of
the callosities of the oscrich. I:luadc. hot ~:
that when the foung osuicb hucbes' Pcwc ts;l
number of callosities already ~ Thcx a!-
losities are ,inntM on the brcut, the pubic: rc:gioo,
and on each ankle-join • 'ow, the c:aJlositia on
breast and pumc region are funaionaJ, fa< owido
resU its body on them when - aouches. &t the
callosi ics on the ankJe.;mnu arc not UKd, or
legs til inwards and so the callooities do '"" bear
on the ground, Instead, the weight of the body 'rcslS
upon an adjaant inner projcaiou of the ankle, and
the skin and scales here gradually thickm and form
an accessory pad',
The interest of this casc Iics in the following fam.
All these caJlosi'cs are "inherited', i. c. intcrn2l
factor, controlling their formation are tnllsm;urd,
but whereas in some (the aeeessory pads) this fl)l"'lm-
tion is evoked by external factors afta' hatching in
each generation, in the others (breast, pubic, and
ankle-joint callosities) these particuJaramrul&aors
,
are not requlred so that t he <lormauon
'w<spbcc
• th"" 'dena for the
before hatching. But 15 IS nO evt
assertion that the internal factors which control the
, f tallosil)' are the
development of the atter type 0
I
 ·\ ELtl:K.\ I IU;,\

e sction <he atun.1 fa 'ors (friction

P ::s:ur<:)~on the body of the an estorl and ~e
inforrnnion as to how the internal
..... no ·.L f
h .arose in thi UtVl in wa( 0 the ttth of

XI'"
P.~EDOt-IORPHO_1 A?\D
GERO:,\,O;\IORPHO I
H E Fe; of e las few chap as bas bc<n '0
T
m;
<ha etch of <he <hcof"ticaJly
hdmx:hronr in produ ing phylogenr has
ible

:ICt1D. ,. in e ",,,Iution of differcn'aninul

e . Ie excepOon of h~-permorpbosis.
- .
..in con Iu..<ion IJUl" th=fof" be dn....n:
I . lu:aJj"ta .,-e c\"OlutionU)" no,-dties an and do
appear a all es in ont~n~') and not
ch- in e .dul .
"
••• as can an do change <he time and
onlcr of eir 'pponnce in the ont<>geny of
e descendant as compared ,..jth th., of the
an or.
3· UlJ.ti':ati·... dilfCf<'DCCS be,.ccn chanc'e,..,
=ill . in hduocbrony, pby • part in
yIogmy in addition to the: introduction of
qualitati.-e DO\"dtie:s.
II iJ It k ...1 l4.l tM.u aubcrifllS rnui. 'fuJIJ
Wtr t4. u/vriuJ 1M" , • ..-_. alllnl 1M
 P.IEDOMORPHOSIS AND
9':l . trw produced
geneuc en l
by some cases of accelera ion
• •
that of the: heart In the chick, or of the
,
,u ch as "rh _a"
oducts in <he chordates. e enect on phylo-
J m
coeo uld seem to be very In . d'Ireet.
geny wo . f
This table has no pretensions 0 completeness,
being composed merely of a few examples selected
at random. On considerIng Jt, however, It does
appear that the. largest ~hylogenetic cffects arc pro-
duced in association with caenogeneslS, devlatJOn,
and neoteny, which are to be regarded as having
pla}'cd a part in the evolution of Amniota, Gastro-
poda, Insects, and Chordates. Caenogenesis, devia-
tion, and neoteny also produce small phylogene ic
chan os, as do adult variation, hypermorphosis, and
acceler.u..ion. But it seems that a big change in
"'olution is mort likely to occur if ancestral youthful
characters becnme those of adult descendan ,than
if ancestral adult characters become youthful char-
acters of the descendan . Caenogenesis, deviation,
and neoteny may then be grouped together since they
are concerned with characters which appear early in
the ontogenr, and they may be contrasted with adult
varia ion, hypermorphosis, and acceleration, which
arc cnnccrned with characters which appear late in
ontogeny. The production of ph)'logenetic change
h)' the Introduction in 0 the adult descendant of
chara ers which were youthful (caenogenetic) in the
:inees or, by means of deviation and neoteny, may
be termed ,o,d.morph.,i, Ph '10 . enetic chan "e..as"'-_ _.....__
C tRO. 'TOMORPHOSIS
a result of modification of ch.a.racttn wh ljl
already adult, by means of adul v . . k.h Wue
morphosi~, and accelcratio~ lmy be~, hyper.
L_ .
morprwJIS, -'''1lQJ l"'''k.-
I f it be true ,hat big evolutioOUJ' chan co, >ad
therefore he large groups of the animal -n
are due to pacdomorphosis, it lhould be poolibl< ..:
corre1~te the characteristics of brge groups "'ith the
essential feature of pacdomorphosis, viz. the appeu-
ance of characur~ in early suges of the ontogeny
of the ancestor. :Sow, the large groups of the anirml
kingdom such as phrla and classes have ","0 charac-
teristics. In the fiTS pbc:c., the anim2ls in anr phylum
or class differ cnnsiderably from animals in otoo
phyla or classes, i. e_ there has been cnnsidenblc
phylogenetic struceural change between <he diJfuCltt
phyla and c/asses. ~en, <he large group<, ph)1a or
classes, contain more numerous different r,-pts
within them han do the smaller gToups luch :u
families, genera, or species. This m.ean~ dl~t the
changes which gave rise to he large group< have
enabled the animals in these group< to gn on evolvmg
further and in more numeroUS directions ~ the
animals in the small group<. The po«~tiahty of
evolving further is called !"'Slid/], ~ "" t bccnhle""'f
ha - .... ~hos .. IS capa 0
necessary 0 show t pilCUU. t'
U
.,,, 'thou
P roduci~ large changes in phylogeny ..,. t
. . d hat gerontomorphos~ IS
sacrificing plasticity, an
not capahle of this.
 P\fDO~IORPllOSIS AND
.' '''
'IT:ir, .:S
.' [() the amount of structural change
ted \l·t know th:tt the adult form is the result of
from forms
C. ERONTOMORPHOS1S

. like larval Echinoderms . 'Ioh e auult ,9J

jJ,'
(flet: ' I I' chordate differs as much from the adult EChInoderm
O

_e•• " and :llso that the C:lr It".T :I. character al'-
ton ! ; . l · as the latter does from the larval Echinodeem,
O .
pt':ltS in onrogtnr the lo~ger It h:ls to proceed with B~t neoteny does not only contribute to the pr.,
irs J('\'tlormenr. Experiments on the gypsy moth ducuon of large structural change; it is also the
hl\'t: pro\-eJ (h:lt the earlier the ma!e characters cause of the retention of plasticity. Experiments on
2ip~1r in y; uld-Ix females, the gre:ltcr IS the change regeneration and tissue-culture have shown that there
that resulrs in the aduh (orm, viz. intersexuality. exists an antagonism between thedegreeofspecialil.a.
Th,u the degree of divergence should be greater the tion of a tissue and its power to grow and become
t"ulier the di\'ergence sets in, is only another way of specialized in other directions. At the outset of
expressing \"on Baa's bw of the greater resemblance ontogeny, all the cells to which the fertiliz.ed egg
bery;«n young forms) and we have seen that in has given rise are more or less similar, and devoid
phylogeny this has been brought about by deviation. of any speci:1lization or differentiation, Such cells
On the other h:\Ild, characters which appear late in are described as being in the embryonic or undifferen·
ontogt-n)' have not much time :1S :1 rule to produce tiated condition. Experiments have shown that
much dr~ct b~fore development ceases. The large embryonic cells are capable of rapid growth, and
:lnlount of structural change eflected in evolution by that their eventual fate is not yet irrevocably deter-
mined. A cell which would normally have undergone
p:ledomorphosis is therefore based on the principle
specialization to form part of the skin for instance,
of deviation, as is illustrated by the supposed evolu-
can be m:1de to turn into part of muscle, or stomach,
tion of the whole class Gastropoda as a result of a
or nervouS s)·stem , as Mangold has shown: As
nri:ltion in the roung.
development proceeds, however, the cells an~ tiSSUes
. Deviation and neoteny combined also playa part , ' I
become Irreverslb y committe
° '0
d specialization and
In ~roducing large structural changes. In anim:1IS in . . " h i ' ~ own line and can nO
dIfferentiatIon eac a ong It:! , d
which the young Stages have to undergo considerable . . fa I n other wor s,
longer alter thetr prospective teo . h
change before the adult form is reached a retention . h' h characterizes t e
of the, form of the roung stage will prod~ce a phylo-
the plasticity of the tissues w IC h Idee
teo
I '\ostat
young stages of deve opmen.t IS I' of changes
genetIC ~~ge as big as (but in opposite direction ° h ,I en)' IS the resU
~o) the difference between roung :1nd adult. This is stages. Now, smce P } og. . . 'bl to expect
. . t IS unpoSSI e
dluSlr:lted by ,h e supposed o in successive ontogellleS, I
evolutIon ofdchor ares
 p.'\EDOl\'!ORPHOSIS AND
9' "' - os co the amount 0 f struccur:d change
I'.r)[, . from forms like larval Echinoderms 'lh
· -~ w'e know that the adult I.form IS the result of
~ nec[cu,
anragen),, and also that the ear. Icr a character ap-
pears in ontogeny the longer It has to proceed with
its development. Experiments on the gypsy moth
have proved that the earlier the ma~e characters
o pear in would·be females, the greater IS the change
P I r ..
chat results in the adu t Jarm, VIZ. lIltersexuality.
That the degree of divergence should be greater the
earlier the divergence sets ill, is only another way of
expressing von Baer's law of the greater resemblance
between young forms, and we have seen that in
phylogeny this has been brought about by deviation.
On the mher hand, characters which appear late in
ontogen)" have not much time as a rule to produce
much effect before development ceases. The large
amount of structural change effected in evolution by
paedomorphosis is therefore based on the principle
of deviation, as is illustrated by the supposed evolu-
tion of the whole class Gastropoda as a result of a
variation in the young.
Deviation and neoteny combined also playa part
in ~roducing large structural changes. In animals in
which the young stages have to undergo considerable
change before the adult form is reached) a retention
of th~ form of the young stage will produce a phylo-
genetIc ~hange as big as (but in opposite direction
~o) the difference between young and adult. This is
illustrated by the supposed evolution of chordates
G ERONTOMORPHOSIS
9l
d .ff' .
chordate I ers as much from the adult Echinod~fm
e al1ult
as the latter does from the larval Echinod"...rm.
B~t neoteny does not only contributt: to the pro-
duction of large structural change; it is also the
cause of the retention of plasticit),. Experiments on
regeneration and tissue-culture have shown that the.re
exists an antagonism between thedegreeofspecialiu_
rion of a tissue and its power to grow and become
spccializ.ed in other directions. At the outset of
ontogeny, all the cells to which the fertilized egg
has given rise are more or less similar, and devoid
of any specialization or differentiation. Such ceUs
:lre described as being in the tmbryo"j( or undifferen-
tiated condition. Experiments have shown that
embf}'onic cells are capable of rapid growth. and
that their eventual fate is not yet irre\'OClbl}' drtrr·
mined. A cell which would normally have undergone
specialization to form part of the skin for instancr,
can be made to turn into part of muscle1 or stomach)
or nervous s),stem, as Mangold has shown: As
development proceeds) however , the ce~ls .and. tIssues
become irreversibly committed to spcclahz.atlon and
dl·ff·erentlatlon
. . eac h a Iong I
·ts own ' line d and can no
. . f t In other wor 51
longer alter thetr prospeCtlve a e. .
the plasticity of the tissues which c.haractcflzes1dth,
. lost at the a rr
I
)'oung stages of deve Iopment. ISheresuto 1 f h go,
C:II1 ..
stages. lOW) since ph)' og~n): IS.' 'bl to exp'"
. . . S It IS UllllOSS1 e
III successive ontog cllIc ,
 ~ Pj\EDO~IORPHOSIS ANI)
J GERONTOMORPHOSIS
much lI11er.lrion ro rake place when the animal has
,;
rtached the larer stages of its development. This is were t he on 1y possible method of I'
why geronromorphosis C1~ only res~lt in the prOduc_ }- acc 'C 5 l eory a r recapitUlation .eVOutlon
I k I' h
Id '
as
1 Id "'ou suggest
tion of small groups of ammals, whIch become mare ph)" ogeny wou gradually slow down d L •
. T an Ult'COrne
and more specialized and incapable of evolving statlonai}', he race would not be able t 1
. a evo ve an)'
furthl"r. But paedomorphosis acts on the }'Oung further,, and
. would be In a condition to WIC h' h te h
stages of development, and if by neoteny the more term. raCial senescence' has been applied. It would
or less embryonic condition of the tissues at these be dIfficult to see h~w evolution "'"as able to produce
young stages is preserved into the adult, these tissues as ~uch phylo.genetlc change in the animal kingdom
as It has, and It would lead to the dismal conclusion
will still be capable of undergoing a considerable
that the evolutionary dock is running down. 1n fact t
degree of further alteration. So, when neoteny had
such a state of affairs would present a dilemma
I acted on the Echinoderm larva, it produced an
analogous to that which follows from the view that
animal which in its adult state was still plastic and in the universe energy is always degraded. If this
capable of evolving further, by both paedomorphosis were true, we should have to conclude that the uni·
and gerontomorphosis, to produce all the diA-erent verse had been wound up once and that its store of
types of chordate animals. The same conclusion can free energy was irremediably becoming exhausted.
be applied [0 rhe evolution of the highest forms of \Ve do not know how energ)" is built up again in
inverrebr:ues, the insects. (It is to be noted that the physical universe although it must happen some·
caenogenesis may also give rise to highly specialized how; but the analogous process in the domain .of
products in the way of routhful forms, but these will organic evolution would seem to be paedomo~phoSlS.
also have lost their plasticity; and as it is unlikely A race may become rejuvenated b)' pushing t~e
that their characters will ever become adult ch:trac- adult stage of its individuals off from the e~d of the.lr
ters, the question of their eA-'eets on phylogeny br . and sue h a race rna}. then radIate out
ontogellles, . In
means of paedomorphosis does not arise.) all directions by specializing an}' of the stages 10 the
. . d"d I til racial s('nesCl:nce
"'e see, then, that evolution by gerontomorphosis ontogenies of Its 10 IVl ua s un . . h .
h · t in again. It 1St 0\\-
produces relatively small changes which sacrifice the due to gerontomorp OSIS se s .. I .
'f has become eXCe5SI\e}
power of changing further J and that on the other ever to be noted that I a race f the
> .' d h younger stages 0
band paedomorphosis produces large changes which over-speClahze .J e~en. t. e a ' ha\'e lost their
(to not sacrifice that power. If gerontomorphosis ontogenies of Its IIldlvlduals m }
 PAEDOMORPIJOSIS AND
""pi.. ..
(IClty.
in this way, .
excessive
. .•
gerontomorph_·
v"l.s
revent the possibility of paedomorphosis.
m.y p . h ,. II .
When Conklin writes t at In every we -tried path
of evolution progress h:u practically come to an end',
he expresses the results of geront~morphosis. And
jf we look round to try to see which of the srecies
of animals living to--day are the anceSlors of the lead_
ing evolutionary novelties of say ten million ycars
hence, focusing our attention only on adult forms
we find it impossible to point out any such ancestor.
If any new orders, classes, or phyla of animals arc
to arise by evolution in the future, it is to the early
suges of animals alive to-day that we must look for
their characterizations.
We may, then, picture Ihe evolution of a race H
a series of revoJutionsj periods of gcrontomorphos:s
alternating with bouts of paedomorphosis. These
terms, therefore) express not only the stage in the life-
history of an animal with which they arc concerned,
but they also convey the meaning of racial senescence
and rejuvenescence. It is interesting to note that as
a result of considerations based 011 a different line
o~ thought, Child has been led to express similar
v~ews. 'I f evolution is in some degree :l secular
ddre~e.n.tiation and senescence of protoplasm, the
, POSSibility of evolutionary rejuvenescence must not be
overlooked. Perhaps the relatively rapid rise and in-
crcue of Certain forms here and there in the course of
evolution may bctheexpression ofchangc50f this sort.'
G E RONTOt\l ORPHOSIS
The conception of paedomorphos· d '11
• IS ren eTl another
service to the" study of phylogeny . If two anllluis
. .
III
a phy 1ogenetlc senes are•
comtv.lred
1'-
~_.
as regarI,Q their
adult structure, one .ahlma! may be •'egar dod as ante_
cedent to the other In that it either d~ ~ not possess
or possesses only partially developed. character
whi~h t~e other animal possesses fully develo~;
I• or, It stili possesses a structure which the other
animal has lost. The antecedent animal in such a
case is called primitive} and the other, Jpecialized or
secondary. These words can only be used to describe
members of a phylogenetic series, and one member
in terms of another, for an animal which is primitive
in respect of one may be specialized in respect of
another animal. For instance, a reptile is more
primitive than a mammal because the reptile's heart
has only three chambers to the mammal's four. But
a reptile is more specialized than a fish because the
heart of the latter has only two chambers.
TOW, it appears to be the case that the str,uctural
changes which take place in evolution are Irr.ever.
sible. 0 case is known in which a race of anImals
after having lost a character acquires thal identi(al
character again. The phylogenetic history of the
· d loss of all
horses has involved the re dueuon an .
fingers and toes except the Jrd. I~ ~me very
. h ·11 b de It With III the next.
exceptional cases (whlC WI e a L--.d
fi ers or toes uol C$
chapter) horses may have extr~ ng 'bT . evolu-
the Jrd, but the principle of Irrever51 I Ity 10
.... 0
 98
p.\EDOMORPHOSIS AND
"" "
I
" akes jt impossible to Imagine that any des GERONTOMORPHOSlS
tIon In . C~n.
( _Le horses may reacqUire. all five fingers and class succeeds another so the h" h ...
d ants 0 UI , Ig er members of
d evolve further on those hnes. The horses the . lower
. .
class may disap~r r-- ,
the 1O\\er
" ones re-
to<S an " I" afe mammg. It . IS from the latter alone, the more un-
comnn"[ted to their phylogenetll.:
. inC and cannot re_ . '
differentiated species of the older class, that t he new
acqUJOre the internal factors
. .which they. have lost . AS
Dolla has put it, the past IS mdestructlble. But while class type can . ..arise.' And the reaSOn wh y th esc
a character once lost is lost for ever, a substitute lower or pnmltlvc members of the class were more
undifferentiated is simply that they have retained the
character may appear which fulfils the same function
plasticity which their previous paedomorphosis gave
as me old character, but is always structurally dis-
them, and have not undergone extensive geronto--
tinct and easily recognizable. As an example we may morphosis) or that they have reacquired plasticity
take the tendency which is observable in the phylt>- by a fresh bout of paedomorphosis, if the previous
genf of the colonial hfdroids for the originally free. gerontornorphosis was not too excessive. For the
swimming medusa [Q become more and more re- same reason) highly specialized products of caeno--
duced to the condition of a sac which never becomes genesis cannot be expected to contribute material
detached and therefore has lost its freedom. In some to be acted upon by paedomorphosis.
cases, however (e. g. in Dicor}'ne), such a sac may It is probable that natural selection was not in-
become detached and regain its mobility, but it can· active in favouring the evolution of groups by paedo-
not be mistaken for a medusa. The muscles which morphosis. The results of gerontomorphosis are
progressive specialization of the adult stru~ure,
served for the locomotion of the medusa have gone
usually in adaptation to some more or .Iess ~estncted
and are not replaced) and for ics locomotion the sac "C Such adaptation entails either the
I
rno d e 0 f lie. . h d"
has to have recourse to a different system altogether, of characters which an leap
cleveIopment or loss . If
viz. cilia. Thus, while structural reversibility appears .' lk of life other than Its own. ,
never to have taken place in evolution) functional the animal 10 any wa I factors arise or change
:eturn to a previous condition using orher inJrrumetllJ then, dimati.c or othe::~~:~n:he animal being unable
IS not uncommon. upsetting thiS mod'~l h e no alternative to extinction.
to evolve further WI av committed to any
" The result of these considerations is to show that . .. imals are not I
But pnmtttve an. t d mod eof \"f Ie.. they are genera -
It IS not possible to derive one animal in phylogeny particularly restnc e . I' d and they will be
from another if the latter is too specialized. CroW has " " d of being speCla \ze )
rzed IOstea
expressed this in the following words: 'As one higher
 PAEDOI\fORPHOSIS AND
,00
j'k j'loundergoextinction3,sarcsuICofcnviron
Jess,"C) ..•
mcntJI change for they ~~s~ss plastiCity. And we
have juse seen chat the prlm~t~vc nat~re ?f an animal
is associated with the plastlclt)' which IS the result
of paedomorphosis.
Ie is wortb noticing that man, whose phylogeny
we have setn (0 be characterized by pacdomorphosis)
largely owes his success to the fact that he is not
ad3pred to any particularly restricted mode of life
at all. Instead, he is fitted for all sorts of habits,
climates, and circumstances. Man himself is general-
ized, not spe<:ialized, and as Elliot Smith has em-
phasized, his body has retained a brgc number of
primitive features which other mammals have lost.
Lastly, a word may be said in connexion with the
bcaring of pacdomorphosis on classification. \,Ve
have seen th:l.t C:l.rly phylogenetic st:l.gcs arc c:l.lIcd
primitive, late oncs specialized or second:l.ry. Y.lc
may call the early ontogenetic st:l.gcs (embryonic or
larval) young, and the late ones adult. Now, if the
~heory of recapitulation were of universal application,
It would follow that an animal which in its adult
stage possesses characters which are present only in
the young stages of development in other animals,
,~'ould always be regarded as phylogenetically primi-
,
tive. But this would lead to absurd conclusions, as,
'
or Instance, that the human stock gave rise to apes,
or that. Perennibranchiatcs gave rise to the other
newts, lIlstcad of vice versa.
GERONTOMORPHOSIS 101
The possession of embryonic or larval charact~rs
in the ad~llt ~oe~. Ilot necessarily prove that the
possessor IS pnmltlve, for it is equally if not more
likely to be due to neoteny and th~refore to be
phylogenetically secondary. Lowe has ~vealed the
most interesting fact that the plumage of the ostrich
rC0l3ins throughout life in the condition of the down
plumage of the chicks of Bying birds. Since the
wings of the ostrich and of the 5O-Cllled Rightl~,
birds must have been derived by degeneration from
those of flying birds, it is to be concluded that th~e
embryonic and larval features of the ostrich have heen
secondarily prolonged and retarded by ncoteny, in
the evolution of flightless birds from flying birds.
XV
REPETITION
E have now come ncarly to the end of our
W review of the relations which ontogeny and
phylogeny bear to one allot hcr, and we have seell
that the Ch:U3cters which appeared in the ontogen Y
I in the ontogeny 0 r
of the ancestor tend to reappear. f
. espondlllg st:lge 0
the descendant, elth~r at a c~;r \Ve have also scen
development, or earher, or lat . f har.ictcr in the
that the accelerated appearance 0 ~ ~onstrueJ :"IS the
ontogeny of a desccnd3lH CI311,notdull ancestral stage
, b k f a COlllp etc a
prcssll1g ac' 0 • I There is then nO
into earlier stages of devc10plllen .
 ,., REPETITION
. . REPETITION Ie]
, Jation in the Haeckelian scnse of accelerated
recapl[U . ' .. the question which at l35t arises is, why are characten
repetition of adult stages.. But there IS repetition.
repeated? The answer is no new one, but it has
Characters which appeared In the ancestor do tend to recently been most aptly formulated by Morgan and
pe ar in the descendant, and, whatever stage these Broman. Characters are repeated because the internal
'P
characters may 'represent,• It
' ,IS d'/li I '
I eu t to aVOid the factors or genes which controlled their appearance in
impression that the enthu~iasm ,of the recapitula_ the ancestor have been transmitted to the descen-
tionist5 is really based on thiS obVIOUS fact of repeti- dants. 'Repetition is, therefore, evidence of affi,,;tJ
tion. If only the recapitulationisls would abandon between ancestor and descendant, which might not
the assertion that that which is repeated i~ the always be obvious by a comparison between the
aduIJ condition of the ancestor, there would be no structures of the adult forms, and this is the real value
re1SOn to disagree with them. Vle have secn that of embryology in the interpreution of evolution.
that which is repeated in the ontogeny of the Given the possibility of heterochrony it is idle to
descendant may represent the cmhryonic or larval assume that the repeated character represents an
just as well as the adult characters of the ancestor, ancestral adult rather than an ancestral youthful con-
and that the retarded repetition of youthful ancestral dition, and therefore any attempt to reconstru<:t the
stages is of particular importance because of the part ancestral adult structure from the embryol1lc or
larval structure of the descendant must be unsound.
which it plays in paedomorphosis. The appearance
All that can be said is that since the ancest~r .g.ave
of characters in the early stages of development is
rise to the descendant it must have been pnmltlve,
caenogenesis, and thesc characters which loom so
and primitive animals are not specialil.ed. Th;.r;for~
largely in neoteny and deviation arc flies in the " b ble that the ancestral adult form I no
Jlaeckelian ointment of recapitulation, for his theory It IS pro a hf I f "
h from the ancestral yout u orm.
W;lJ bound to treat them as exceptions to his rule depart as muc descendant departs from Its
of evolution. It is because these early developed the adult form of the lraJ adult structure
r Th s the anCes
c~araeters are not exceptions in phylogeny, together youthful Iorm. u d
can only be inferred an not pr
oved from a study
~lth the fact that phylogeny is the result of ontogeny
lI1stead of be'Ing .Its cause, that we reject
. k J' 5 of embryology. ., of characters in suc·
Haec.c The nature of the repeutl~n esting light on the
theory of recapilulation. . h OW! an mter
We. .are Jef t Wit'h heterochrony playing upon t h e cessive ontogenies t r II included under the
·ch are U5ua y
phenomena W hI
repetition of characters i .suc Lv.e..ontoD.oenciJ'OSo-".ll!d!.... l_
 REPETITIO~
.'"
""'" ....ri=•. In the borse :ill the linger.; and to<s REPETlTIO~
'0\
ffb<m lost etteptthe "rd on eoeb limb. Wben,as cessn of ~mcnt'... "on and of fomu.tion of
. e
..,.,. ~ &.pp<n a borse b:>s an extra ling.. reproductl\~ gb..nd resp«ti,-th·. XonnallT the:
• io addition the Jed J l [ presents :1 eatain btter are quick enough to ~ e ~~ai\
or roe ce to :l IDOf1: primiti'"e pbylogenetic condi_ gbnd before the processes of ~entationcan di . e
.~
_ ..n wh.'eb the reduerion of .finger.;
th and __'oesbad
, 00, it up between se\'ttal ~en But if these Ff'O.
adtd 2S f.u 2S cy h:l\'e In· e nOUllal structure cessesa.rerdati·..dydda~~theproct::sso.of ... n-
pioc _L'_ tatioo will effect 11 partitioning the: ~roducti''''C
of the OlOdern horse. But oes "'"' mean that the
gbnd before the latta is deiinitdy formtd) and so
~:~bo<.e with = linger.; b3S 'gone bad;' tn '~produce' the ~cestral condition. AtuisQ .
0:; . . . .<stul type: Gegenbaur showed some time ago
therefore) due to the l"'Cproduetion of 2 Stt of i.
<!:a.: this could be aliirmed, an the =ent exten-- tions (a definite system of raetioo-spttds) .hich
sioos "our kncn<iedge of be=litJ have provided an obtained in the ancestor.
~ of ese cases of so-called at:l\;sm. Analogous resul of an 2tl'--is .c D2.~ hl,''''C bttn
In tM men primitiYe insects such as Iap}'"X or obnined br RunnstrOm in the c:ourse of his ~''''CS . g;t-
. the rtproducti,·c organs are repeated in tions of ~rimenral embryo1og) of ~erms.
eodl of seven segments of the abdomeo. Tbat this The l-an...e of Echinoderms art' dwaeteriz.ed ~a.
is a pri:cime fealUIe cannot be doubted, for the all b\' the poss'"SSion of three p:airs of bodr-a.n~.
C'SStttt e segmentation of the bod)' is the b utr.10~ th e bn.. of Antedon the 1Stc.and. ed :nd ca"bes
If. bo_
tepeci<ian of the rq>rodu 'ye glands, as in many th . ht side are not nornully ,oem . • -
on e ng _. twO b\' a tnn5-
'u ws. In the higber insects, of ,..bieb the gyps)" e,,-er a bn-a of Antedon 15 cut In '. _
• . b2lf dr\"t'lops thoe ca"n~
is an cnmple, the reproductive glands are ,·erse cut) the postenor Anttdon bn-a and
IocaEz.cd in one UKio:ninll segment on If 2Jld are not thereb)" differing from a. n<>rnWEchinoderms. Tbis
" ....ned. Goldschmidt b:>s found, in the course of resembling.the.1U'\"2.e of ~:ed bv the r«StJ.blish-
C1iQi-e.ntal breeding of races which differ in the case of aCl'\'1sm IS to be up di .' brought about
tral set of con bons
rdativ< speeds of developmen., that gj-psy moths can ment of an ances inhibiting factOr which \OS
be obeained in ...bieb the reproduetive glands are by the remo,-a) of an . half of the 11[\"2 was
. ed b the anteOO r
~ r repeated4 This condition, which is elimmat v: en . -0'cspnM.nnces an
be of other au":> r·-----
'0
termed hptmul], is be explained b)" the different cut off. A num r ,
be produced in the same v;2.).
rate::s of actions of the factors wbich control the pro-
- r
 ~EPETITI N
,06 ,
J tly we may consider some atavistic phenomena '"7
m. m:an.
..as 'It was
• shown . in Chapter VI J J that the
evolution of man was In a large ~ea5ure due to XVI
"ny and retardation) and we saw In Chapter III CONCLUSIO:"S
nCO . J'
rhat the rate of development In mamma 5 IS under ,
Ihc control of the ductless glands which secrete
.., hormones. During human phylogeny, therefore, the
T ilE !ine of argument which form' the iuhjet t
of thiS essay may be lummariz.ed u follo'A'i:
Ontogeny is the result of the actIon of external
ntc of development of the body must have been factors in evoking respon5Ct from the inteTnal facton
progressively retarded by the action of the ductless of an animal to which the latter were t",n,mitled by
glands. Suppose now that for some pathological inheritance from ill parents.
reason one or other of the ductless glands ceases to Phylogeny is a series of adult forml which are
exert this retarding function, the result will be the disconnected and causally unrelated to one another;
production of a man with so-called 'pithecoid' char- each adult form being the result of an ontogeny
acters: the hair will be over-dcvcloped, or the skin I which differs from the previous one.
dark, or the brow-ridges accentuated, or the jaws Successive ontogenies are rdated to one another
larger, or the length of the bones will be increased, by the transmission of internal factors from fertiliud
or sexuality will be precocious, or the sutures between egg to fertilized egg. .
the bones of the skull will close up prematurely. Modifications in ontogeny (in a constant environ·
These features are atavistic in that they result from ment) arc due to changes in the intern~1 facton.
a reversion to the set of circumstances under which Phylogeny is therefore due to m~lfied onl~e~y.
development took place in the ancestor. Phylogeny plays no causal part III determining
Atavism is, therefore, based on an inheritance of ontogeny. .
The internal factors exert their effects at certain
similar internal factors by the descendant from the
ancestor, but, like normal development, atavism is definite rates. .
Modification of the rate of action of th~ Internal
a~ualJy brought about by that new creation of an . . .es will result In hetero-
I factors III succesSive ontogem
aOlmal, which constitutes ontogeny. The atavistic
feature is not directly inherited from the ancestor as chrony. ... f
· . b ought about by acquISItion 0
su~hl for nothing is solely inherited without also I
E vo ullon IS r ad . f novel
being acquired.
'd
qualitative nove Itles, an by the pr ucuon 0
"
108 CONCLUSIONS
• CONCLUSIONS
situations by quantitative alteration of the ratc of
Atavism is due to the bl" 1°9
action of the internal factors. geny of the desce d re.efsta Ishmcnt in theOllto-
New characters may appear at all stages of onto- n ant 0 a set of .
which was present in the 0 t f CIrCUmstances
geny, and by heterochrony they may be retarded or I . n ogeny 0 the ancestor
accelerated, so as to appear later or earlier in subse- , t goes without saying that even if Ih .
fo th h
• r 1 ere " are correct ' they d0 not prOVIde .
.
e views set
an
quent ontogenies. exp anatlOn of evolution , for th ere remainS . the
Characters present in the carly stages of ontogeny
problem as to how and why novelties arise, and why I
have (provided that they arc not too specialized) , heterochrony acts upon them '·n those cases In
. W h·Ie h
.
plared an important part in evolution by paedo- It does. But it is claimed that after dethroning the
morphosis, resulting in large structural changes theory of recapitulation, we are able to make a beuer
without loss of plasticity. synth~sis of our knowledge of embryology and
Characters present in the late stages of ontogeny evolution.
have plared an important part in evolution by gcr-
ontomorphosis, resulting in relatively small structural I XVll
change with Joss of plasticity. BIBLIOGRAPHY
Paedomorphosis and Gerontomorphosis act inter-
mittently in thephylogenyofa race, the former produc- , ACASSll. L. An ES$;IY on Classification. London, 1859. p. 269·
AIlISTOTLE, in Slllen. C. G ree\.: Biology and G reck Medicine.
ing racial rejuvenescence, the latter racial senescence. Oxford, t922, p. 45·
BAtIl. K. E. \'0", 'Ueber EntwicklunVgeschichte dcr Thitrc.'
Recapitulation, i. e. the pressing back of. adult Beobachtung und Rcflelion. Konigsbcrg, t828,
ancestral stages into early stages of development of BATESOS, W. Problems or GenetiC1. Yale, 19'3. p. 61·
descendants, does not take place, although there may Bou. L. Das Problem der r.lenschwerdung. Jena, 19 z6 .
BoSSET. C. ContempLation de La nature. AmsterJam, 11~'
be acceleration of one structure or another. BII.OMAS. I. D1S sogenannte Biogenetische Grund~tz. Milnchtn
Repetition of ancestral ontogenetic stages in the und Wi~luden, 19 l0 •
ontogeny of the descendant) whether retarded or CHAMI'Y. C. Se:rualileel hormones. Paris, 19 l -j..
1
CHILO. C. M. Senescenct' and Reju\'enescen«. Chiaga, 19 }.
accelerated) is due to the transmission of internal
I p. 46+. . 1 \" 1
factors from ancestor to descendant. _ Physiological Foundalions of Behavior. New, or, 19 +,
Similarity in ontogeny b etween any anima . 1· • •• E G Hertdit, and Em·ironment. Princeton, 19!!'
s IS a CONJ(U.,. • • 'J I r
Caow. W. B. 'I'hylogeny and Ihe Natural Spttm. ourna 0
proof of their affinity, and no evidence as to the adult
GenctiC1, 17, 1926. p. 12 9,
structure of the ancestor.

\
 BIBLIOGRAPHY
"0
. C. The Origin of S"....ies. (Edition: London, 19") • BIBLIOGRAPHY
DA.~'I1'i, r--
GOODIllCH, E. S. 'Melameric ~ " III
p. 61 .. . . . .
lerIy Joumal of Microscopical ~
ljw('Jltl.tlOl\ and H.-...t- , f\..
-,,-. ),....u·
DtTWu.u. S. R. 'uperimentaJ Studies on Morphogenesis in the
- - Living Organisms Oxford 6,59,1913, r· u,:,.
Nuvous Srstem.' Quarterly Review of Biology, I. 1926, p. 61. ' . ·'9 4-.p. I.
Z
H AlCUt., E. Cenerelle ~'1orphokagie der Orp . .
Oouo, L. 'Us Cepbalopodts deroulis ef I' Jrre\'enibilite de 1'£"olu_
~. Bertin,
cion.' Bijd12~ tot de Dierkunde, 22, '921:, p. 21;.
DOllO£S, J. E. 'Methods of E\·~Jution.· Scien~ Progress, 18,
, 1866. 11. pp. 1 & 9.
HU.I.150X,j. W. H. 'The InducrionofMe1anise iu_
and its Evolutionarv Si......a:<:aDCt..
"
I.q:.iJOPlCI'l
'92-4-, p. ;60. -, 6·.... ' . aN~, 119, HJZ~
P· 12 1· •
Eu,u, G. H. T. Organic E"olution. (English translation),
HAJ.VEY, \V. 'Exercitario anatomia de motu cordts et •.
Loodoo, I 90·
, animalibus' in Adriani Spigelii .'\mromica.. ,,_,-,~!,1fII$. .m
EWOTSxITII,G. Essapon the El'olution of Man. Onord,1927. "" .---............ 1 5,
p. W 11.
FO..D, E. B. 'The Inheritance of Dw-atnng in Gammarus che,,·
HuT..... lc. O. 'Ueber die Stellung der \·erg\eicbmd.CIl Enhloic.....
fnl:r:i.' Journal of Generics, 20, 19 28, p. 93· lungslehre zur \'ergleicheoden AlII.tomie, :tor SVltewu.. . und
FOAo, E. B. and Ht:XLU, J. S. 'Mendelian Genes and Rates of Descendenztheorie.' Handbuch der \'erg'lei<:~CIl und ct.
Denlopment in Gammarus chevreu:li.' British Journal of perimentellen Entwicklungslehre der Wirbeltierc, III. It
EXjXrimental Biology, 5, 1927, p. J 12. Jena, 1906, p. 1~9'
j."un. V. 'Ontogenie uod Phylogenie.' AbhandJungen zur , - - 'Das Werden der Organismen.' Jen., 1916, r. 216.
Throrie der organischen Entwid.lung. Berlin, 1927. HIS, W. U nsere Ktlrperform und das phrsiologitthe Problem inrcr
GARSTASC, W. 'Preliminary Note on aNewThcoryofthe Phylogeny Entstehung. Leip7jg, 1814.
of the Chordata.' Zoologischer Anzeiger, 17, 189+, p. 122 & HURST, C, H. 'The RccapiruLnion Theory.' Narur.al Scienct. 2,
Quanerly Joornal of Microscopical Science. 72, 1928, p. 5 I. 18 93, p, '95· ., . . .•
- 'The Theory of Recapitulation. A Critical Restatement of the • HuxLtY. J. S. 'Time Rdanons In Amphibian Mel&morphOilt.
Biogenetic Law.' Joum.al of the Linnean Society of London, Science Progress, 17, 1923, p. 606. ."..
Zoology, 35, 1911, p" 8 I. _ 'Constant Oiffert:ntial Growth-Rates .nd their Slglllfi.:a1'lcc.
- - 'The Origin and Evolution of Larval Forms.' Report of the Nature, 114, 192+. p. 895" .. , .
~ 'Ph)·I_o)·
H YArf, t\. of .n Acquired Cbaraclenmc, Amenan
British Association for I.he Adnneement of Science, Section D, ~e-"

191 Naturalisl,27, 1893.P· 86 5· - .• P ft\fngs

.
GEC£SBAUJ., C. 'Kritische Ikmerkungen nba Polydactylie als , _ 'The Ph,.logeny of .n Acquired CbaraclmtbC. roc I
of the Amerian Phiiolophical Sociel)·. 32. 1~9'f' r· 3:J'
bu
Atavismus.' Morphologisches Jahrbuch, 6, 1880, p. ,84. " C A Three L«ruTd on ~nlrobiotuu. (It
G ILTOI.D. H. The Disorders of Postnatal G rowlh and D('\'elopment. hA.... EU. • • .

London, 1911. S,b)"eelS. Copenhagen. 19 28 . d d· C - " , '

. . h GrundplZ un Ie ... ~"•• " •
GOLDSCHWIDT, R. 'Einige Materiali('n zur Th('(lri(' der abgestimm- K£IB£t., F. 'Oa! Blogeneusc ~ nd Enlwicklun!'geteh~ht~. 1,
Ergebnisse der Ana100lie u
len Reahionsgesch....-indigkeiten.' Archiv fUr Mikroskopische
Anatomie und Enlwid.:1ungsmechanik. 98, 1913, p. 292. 1897,P·111, -aDltomillChe und entwicUunr-
.
KuJ:EJIlTHAL., W. Vergle,chend S' enen Semon's Foe·
- Physiologische Theori(' der Vererbung. Berlin, 1917. . . U rsuchungcn.n Ir .
geschlchtl~che nte" he Denbchriften. 7, ,891. p. H'
COODI.ICH, E. S. 'On the Coelom, Genital Ducts, and Nephridia.' ,chungsrelscn 4. Jenalsc
Quarterly Journal of ?\ticroscopical Science, 37, t 895, p. 4+1,
 1f1 BIBLIOGRAPHY
81 BLIOG RAPHY
1..l,lolA.CK, J. B. P. A. DE 1\1. 01:. Philosopniezoologique. (Edition frondosa G unneru! und P I ha. II J
C. i\lartins), IJ;lriS, 1873, I, p. 2lJ. 1\.1 useum Aarbol 19 18- so us~ nt~pu5 Slr&ucnfdt. 1k'&a\1
LANe, W. D. 'The Evolution of Ammonites.' Proceedings of the RUSI'nROM J '2u . 19· atuTvldenskabdig raekle 5
Geological Association, 30, 19'9, p. 50. , . "r CJipcnmentellen f\nalvu. d £ _ -
Antedon.' ArchivlUr Enty..· II' 1c
,- cr nlv. klung \"01'1
Lowr., P. R. 'Studies and Observations bearing on the Phylogeny of Ie ungsmccha.nik
R USSELL, £ .S. Form and Function l...ond 105, 1
' 19 5op·6J.
the Ostrich and its Allies.' Proceedings of the Zoological Society SEDCWIC". A. 'On the La..... of ~ I oa,19 t 6.
of London, 1928, p. t 85· e opment commonl)- I
\'on Baer's Law, and on the Si,nifica fA nown as
MAcBRIDE, E. W. Text-Book of Embr),ology. I. Invertebrata. - E nee (I nccstr.l Rudi
ment! 10 mbryonic DC\'c\opment' Q 1 -
London, 191+, p. 6;:0. P J'. uartcr y Journal of
Microscopical Science' 36, 189'.,.,.).
~lA~'COLD, O. 'Transplantationsversuche zur Frage der Speufitit SERaU, M. 'Recherches sur l'anuomie com tt d .
und der Bildung der Keimbliuer.' Archiv rur mikroskopische . cb' I Qu pat es anl~UI
Invert. res.: e sont par rapport aUI vcrttbres el aI'homme
Anatomic und Entwicklungsmechanik, 100, 192J, p. 198. Ics anlmaux m\"ertebt61' Annales des &ienccs Naturdlcs,
~lECI(EL, J. F. Beytriige zur Verglcichenden AnalOmie. 2. Ent- J 1,2, 182,,"- p. 2<\.8.
wurf einer Darstellung det zwischen dem Embryozustande der SEWUTZOW, A. . 'Olxr die Beziehungen l.....;scheo dcr Onto.
haheren Thiere und dem permanenten der niederen Stan genese uDd der Phylogenese der Tiere.' JenWchc Ztiachrift
findenden Parallele. Leipzig, ,811. fur Natunvissenschaft, 63,1917, p. 51.
M£IlNUT, E. Biomechanik. Jena, ,898. S~llTIl \VOOOWARO, A. Proceedings of the Linnean Society of
.M £T'SCHNI KOFF, E. 'Embryologic der doppeltfUGigen M yriapoden.' London. I 35th Session, '923, p. 30.
Zeitschrift fOr wissenschaftliche Zoologic, 24, 187+, p. 253. SPATH. L. F. 'The Ammonoidea of the Gault.' ~Iemoin of the
.MO.CAN, T. H. A Critique of the Theory of Evolution. Princeton, PalaeontographicaJ Society, '923, p. 65 .
19 16 . _ _ 'The Ammonites of the Blue Lias.' Proettdings of the G~
- Heredity of Embryonic Characters.' Scientific Monthly, 18, logical Association, 35, '924, p. 186.
19 2 +. p.;. _ _ 'Notes on Yorkshire Ammonites.' The Naturalist, September
- The Theory of the Gene. Yale, '926. 1925, p. 268, and May '926, pp. 137, 139· .
MULLER, H. J. 'Artificial Transmutation of the Gene.' Science, 66, STOCItAJ.D, C. R. 'The InBuenee of Alcohol and oth« Aru.tstheucs
1927, p. 8,... on Embryonic Dcvclopm~nt.' Am~rican Journal of Anatom}",
MDun, Fa,rrz. FOr Darwin. Leipzig, 186+. 10, '910, p. 369'
Tllo)\I'SON, D'A. W. On Growth and Form. Cambridge, '9 17,
NA£F, A. Die individueJle Entwicklung organischer Formen 315
Url:unde inrer Stammesgeschichte. Jena, '9'7. pp. 54-, H9· . 10 d" ' j
-Idealistische Morphologic und Phylogcnctik. Jena, '9t9' T - A'.-,-,-,1 De\'{~lopment.'
1l0MSON,.. .. 1 he Denl" teor, ,19·'
OSBORN, H. F. 'Origin of Single Character Differences.' American p. t '9)- 'Ober die Rlidbildung der Kiemenbogen bei den
Nuuralist, 49, 1915, p. '9J. V EJ.SLU¥S, . . 11 , p. 9-)..
--'
&1;lC hII. B'-J-dn,en tot de Dlerkunde, 22,19
RElcH~"T, C. B. Vergleiehende Entwid:e1ungsgcschichtc des \VEISMANN, A. The Evolution Theory.
(E Ii h tranWuon)
ng s '
Kopfes der naeklen Amphibien. Konigsberg, ,8 J8.
RIDOU, O. 'Internal Secretions in Evolution and Reproduction.' London, 19°4, pp. '7?' 186 die SLlmmesgeschichte der
\VURTI'.MDUG£Il, L. Studlen Ii ber
Scientific Monthly, 26,1928, p. 216. Ammoniten. Leipzig, ,880.
RUNssnihf, J. and S. Obe.r die Entwicklung von Cucumaria
 '.',t-. ".
I J....,,'·'I'
t;:t,~
' ........, ""I

II

101..,', II, At,

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 INDEX

Perif»-tUJI, 44· .kin, colour of, 61.

phylogeny. 5' 2.8. Smith. Wood"ud
. , S.
pluticily, 91. IOmauc mductioo, 1 S.
I'liot.rema. H. Spa~h,. I I, .p, 5 I.
Polyltomum, 17. 57· s~clalhed characters, 9].
Portunion, 4!. Iplnl cleavag~ 41.
primitive chanCU!I"I, 97. Stockard, I).
prlmro5e,2.J.
progeria, 2.6. teeth, 6..., 73.
Proteu,. 2.7, 58. tendon, fibrel of, 16.
Protohipput, JO. Thompson, 19.
J)iiloans, 5 J • Thornton, 61.
Python, 71. thyroid, 14.
titanothcret. 31, 8J.
lUna, 2. 5. Tornaria, 66.
recapitubtion, 6, 79, 101. tonion, GUlropod, p.
reduction, 37, 71. tracheae, 70.
Reichert, So. transmission of facton, 14·
rejuvenete.encr., ncuJ. 95. Trochosphere, 41.
repetition, 101. Tulotoma, p.
ret.atduion, J8. 7S. Typhlomolge. 17. 58.
Riddk.2.5·
Runruu6m, 105. Unio, 41, 4j.

K.ak of heingt, 2.. Vdiger, 49.

IeCOndary charaetcn, 97. Versluya, 55.
Sedgwick, 10, H. von Bacr, J. 45, 56. 7)·
IegmenLllion, 54.
ICnelCCnoe, rilCial. 95. Wcismann, 8.
Serm, J. WQrtcmbcrl::cr, 7. 5 J.
Se"eruow, II, .. 6. 56, 77.
Siu.ri" "', H. XiphophoruJ, H.