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South African Journal of Science Vol.

73 April 1977 121


The binding of both toxins to synaptic substances. or Naja melanoleUL'Q venom. J. bioi. Chern., 247, 2866-2871.
inactivation of synaptic substances involved in transmission. was , ShipoJini. R. A.. Railey, G. S., Edwardson. J. A. and Banks, 8. E. C. (I 973).
Separation and characterization of polypeptides from venom of
fairly stable because the effects were irreversible for a period of Dendroaspis ~'jridjs. Eur. J. Bfochem., 40, 337-344.
at least three hours in both instances. ~ Sato, S., Abe. T. and Tamiya, N. (1970). Rinding of iodinated erabutoxin
b. a sea snake toxin, to the endplates of the mouse diaphragm.
Toxicon. 8, 313-314.
Conclusions 9 Miledi. R .. Molinoff. P. and Potter. L. T. (1971). Isolation of the
cholinergic receptor protein of Torpedo electric tissue. Nature, 229.
Toxin '1. did not prevent primary afferent depolarization and 554 557.
dorsal root potentials. Yet presynaptic inhibition was ineffective I" Chang. C. C. and Lce, C. V. (1963). Isolation of neurotoxins from the

in reducing synaptically elicited motoneuronal responses. Toxin ~ venom of Bungarus multicinctus and their mode of neuromuscular
therefore caused hyperexcitability of frog spinal motoneurones. action. Arch. intern. Pharmacodyn., 144,241-257.
II ue, C. Y. (1972). Chemistry and pharmacology of polypeptide toxins in
presumably by primarily lowering the reaction threshold of the snake venoms. Ann. Re~'. Pharmacol., 12,265-286.
neurones concerned, and not by blocking presynaptic inhibition. Il Szezepaniak. A.C. (1974). Effect of (x·Bungarotoxin and dendroaspis
The hyperexcitability produced by toxintJ, on the other hand, neurotoxins on acetylcholine responses of snail neurones. J. Physioi .•
was apparently mainly accomplished by blocking presynaptic 241, 55P-56P.
11 Roper, S. and Diamond. J. (1969). Does strychnine block. inhibition
inhibition.
postsynaptically? Nature, 233. 1108-1169.
Received November 29. 1976; accepted January 19.1977. I~ Holemans, K. C. and Meij, H. S. (1968). An analysis of some inhibitory
mechanisms in the spinal cord of the frog (Zenopus lan'is). Pfliigers
I Holemans. K. C.. Meij, H. S. and Meyer, B.l, (1966), The existence of a Arch., 303, 287-310.
monosynaptic reflex arc in the spinal cord of the frog, Exptl. Neurol.. I' Grinnell, A. D. (1966). A study of the interaction between motoneurones in
14, 174 186. the frog spinal cord. J. Physiol. Lond., 182.612-648.
Bremer, F. and Kleyntjens, F. (1917). Nouvelles recherches sur Ie I. Holemans. K.C., Meij, H.S .. Meyer. B.l. and Loots. 1.M. (1967).
phenomene de la sommation centrale d'influx nerveux. Arch. intern. The use of the South African frog in the study (If spinal reflex
Physiol., 45, 382-413. physiology. Onderstepoort J. vet. Sci., 34,2619-2632.
Meij, H. S.. Holemans. K. C. and Meyer. 8.1. (1966). Monosynaptic I' Rotes, D. P. (1971). The amino acid sequence of tox.ins tX anCl P from
'transmission from afferents of one segment to motoneurones of other Naja nil'ea venom. and the disulphide bonds of toxin a. J. bioI. Chern.
segments in the spinal cord. Exprf. Neural.. 14,496-505. 246,7383-1391.
Loots. J. M. and Meij, H. S. (1972). Monosynaptic testing of synaptic 18 Rotes. D. P .. Strydom. D. J., Anderson. C. G. and Christensen. P. A.
efficiency and motoneurone excitability. S. Afr. rned. J., 46. 203-207. (1971). PurifLcation and properties of three tox.ins from Naja nivea
Loots, J. M., Meij, H. S. and Meyer, 8. J. (1973). Effects of Naja nivea venom and the amino acid sequence of toxin <5. J. bioi. Chern.,
venom on nerve. cardiac and ~keletal muscle activity of the frog. 246,3132-3139.
Br. J. Pharmacol.. 47, 576-585. 19 Davidoff. R. A. (1972). Gamma·aminobutyric acid antagonism and pre-

Bates. D. P. (1972). The amino acid sequence of toxins band d from synaptic inh.ibition in the frog spinal cord. Science, 175,331-333.
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2010).

Research Letters
frequency of occurrence (68%) of the plankton feeding haJtbeak
On the Life Cycles of the Cestode Hyporhamphys knysnaensis (Smith) in its diet. Thirteen
Ptychobothrlum belones and specimens of H. knysnaensis were caught during seining
operations on Lake St Lucia, and one of these was found to
Nematodes of the Genus Contra- harbour typical pseudophyllid plerocercoids in the epaxial
musculature. Only T. leiurus and the tenpounder, Elops
caecum from Lake St Lucia, machnata (Forskal), feed on H. kynsnaensis to any significant
Zululand extent, but none of the 81 J E. machnata e);:amined had
tapeworms of any description in the intestine. These
During an investigation into the feeding biology of piscivorous
fishes and birds at Lake St Lucia (280 0'S, 32°30'E), a record observations, together with the obvious similarity between the
was kept of cestode and nematode infestations. A collection of scolex of the plerocercoid and that of the adult P. belones,
these parasites was identified as far as possible by Dr S. Prudhoe indicate that H. knysnaensis is a second intermediate host of this
of the British Museum (Natural History). In relating the cestode. H. kn.}'snaensis fultiJs two important requirements for
incidence of infestation to the food preference of their hosts, it being a second intermediate host of a pseudophyllid specific to a
has been possible to identify, for the first time, a second species of T.vlosurus: 1) its narrow cylindrical form makes it a
intermediate host of the cestode P(vchohothrium belones suitable prey species for a predator with a limited gape, such as
(Pseudophyllidea, Bothriocephalidae) and to confirm aspects of the garfish, and 2) its diet includes large numbers of Copepoda,
the life cycle of the nematode Contracaecum spiculigerum among which the first intermediate host is likely to be found. No
(Ascaroidca. Anisakidae), This is the tirst record of these two pleroceroids of any description were found in Gilchristella
species from South Africa. aestuarius, Thr.vssa vitrirostrus. Stolephorus commersonU,
Hilsa kelee and Hepsetia breviceps which, together with H.
Ptychohothrium helolles (Duj.) knysnaensis, constitute the entire plankton feeding fish fauna of
Adult P. helones, attaining a length of 200 mm, were found in Lake St Lucia.
the intestine of the garfish, Tylosurus leiuros (Bleeker), and were P. belones has been recorded from several Tylosurus species. I
present in 29 of the 46 specimens examined. As many as six of Of these the diet of only T. leiurus has been sufficiently studied
these tapeworms may parasitise in a single host. Our material to show a preference for a prey species whose feeding habits
differs from that described by Yamaguti l in having a more make it a suitable intermediate host. We have been able to
broadly spatulate scolex. but the structure of the reproductive examine six specimens of Tylosurus crocodilus (Leseuer) from the
systems appears to be the same. Correlated with the high Kosi Estuary (26°54'S, 32°52'E) but these all had empty
incidence of tapeworm infestation (63%) in T. leiuros is the high stomachs and were free from tapeworm infestation. It would
L22 South African Journal of Science Vol. 73 April 1977

A similar situation appears to obtain in Lake St Lucia,


Table I. Occurrence of Coruracaecum larvae in the mesenteries
Sarotherodon mossamhicus, primarily a benthic feeder. is very
of various fish hosts from Lake St Lucia
likely to ingest the newly hatched infective larvae directly, It
may, in turn. transmit these larvae to the bird final host. or it
Percentage may be eaten by one of the piscivorous fish listed in Table 1. In
Number Conlracaccum the latter case the larvae will find their way to the mesenteries.
Host species Common name Diet examined infestation where they will continue to grow. but will not mature. It is
significant that S, mossambicus is the only fish species common
Sarotherodon mossambicus Mozambique Detritivore/ 231 I :5',~)
(Peters) tilapia omnivore
to the diets of all piscivorous birds and fish of the St Lucia
system. with the notable exception of Tylosurus leiurus. a
Muraenesox bagio Pike Piscivore 13 8(,~J
(Hamilton-Buchanan) conger species unlikely to ingest prey of the shape of S. mossambicus
Lichia amia (L)
and also the only predator which appears to be free of
Leervis Piscivore 20 -'()1I
Contracaecum larvae.
Elops machnotQ Tenpounder Piscivore 81 L 3 f !h
(Forskal) Whether or not the piscivorous fishes serve in any way to
complete the life cycle of the Contracaecum species which infest
Plalyccpko{us RaTtail Piscivore 16 19',~,
indicu$ (L)
them is uncertain. since we have no information on the juveniles
flathead
of these host species, The birds of St Lucia. including pelicans.
A rgyrosoma holo- Kob Piscivore 408 18 f )(,
tend to feed on fish of less than J50 mm standard length, which
lepidotus (Lacepede)
excludes larger predatory fish from their diet. An exception is the
Clarias gariepinus Sharptoolh Piscivorel 28 46f~()

(Burchell)
fish cagle. Haliaetus vocifer (Daudin), which does take larger
catfish omnivore
prey. Faeces and regurgitated pellets of H. vocifer have not been
examined, but records of successful stoops show its diet to
include ] 3% C. gariepinus and 5% E. machnata. The ingestion
seem from the number of final host species and their widely
of larval Contracaeum is therefore almost certain and infestation
separated distribution that there must be several second
possible. However. Thomas] has shown that domestic ducks and
intermediate host species of P. helones. each forming an
fowls are immune from C. spiculigerum infestation. implying a
important dietary component of the genus Tylosurus. H,
degree of host specificity where the final host is concerned. C.
knysnaensis. for example, is endemic to South African waters,
while T. leiurus is an Indo-Pacific species and is therefore likely microcephalum is a common parasite of pelicans, and is found
to feed on other hemirhamphidae in Mozambique and further everywhere pelicans occur (Prudhoe, pers. comm,) and C,
norlh. spiculigerum has only been recorded from the genus
A full description of our material, including the plerocercoid. Phalacrocorax.
will be pubJished when enough plerocercoids have been collected With the St Lucia record, C. spiculigerum is now known to
to permit sectioning. Examination of Copepoda collected from occur in the Nearctic, Palaeractic and Aethiopian regions. This
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2010).

Lake St Lucia is proceeding in an attempt to find the procercoid wide distribution may well be linked with that of cormorants.
Cross-infestation between P. auritus auritus and P. carbo is
stage of P. belones and to identify the first intermediate host.
possible in north-eastern North America, while P. carho has a
Contracaecum spp. continuou:; distribution from North America through Europe
Larvae of various sizes belonging to the nematode genus and Asia and down the length of Africa. Occurrence of C,
Contracaecum were found In the mesenteries of several fish spiculigerum over this range would then reguire only a degree of
species from Lake St Lucia (Table 1). Active living larvae have non-specificity towards intermediate fish hosts. which seems' to
also been obtained from the regurgitated crop contents of the be true for several species of the genus 2 1>: C. aduncum. for
pied kingfisher, Ceryle rudis (L), and the white peLican, example. is known to occur in 40 species of marine and brackish
Pelecanus onocrolalis L, while living adults were found in the water fish.2 The Anisakidae arc primarily parasites of fishes,
faeces of the white-breasted cormorant, Phalacrocorax carbo with only some Anisakinae (including several Contracaecum
(L), The larvae from the white pelican are probably species) and Raphidascarinae parasitising higher vcrtebrates. 4
Contracaecum microcephalum (Rud.) and the adults from the This has probably contributed to the marked adaptability of the
white-breasted cormorant are identifiable as Contraceacum genus Contracaecum to a variety offish intermediate hosts.
spiculigerum (Rud.) (Prudhoe, pers. comm.), a species described This work was carried out while one of us (A. K, W.) was the
as "common in estuarine birds" in Europe2 and also recorded holder of a C.S.I.R. Post-Honours Bursary. Permission from the
from the common cormorant, Phalacrocorax aurilUs auritus Natal Parks. Game and Fish Preservation Board to work at St
(Lesson) in Illinois, U,S,A.' Lucia is also gratefully acknowledged,
ThomasJ has described the life cycle of C. spiculigerum from A. K. WHITFIELD
Illinois: the larvae moult twice within the egg, but do not J.HEEG
exsheath; on hatching they become attached to the substratum, Department of Zoology,
and although able to swim, they show no activity responses to Unil'ersily of Natal.
movement in the water; if eaten by a fish. exsheathment takes Pielermarilzburg.
place in the intestine, and the larvae find their way to the
mesenteries, where they grow but remain immature; maturation
Received January 25. 1977.
takes place once the fish intermediate host is eaten by the final
host. in this case the common cormorant. Natural infestation of ! Yamaguti, S. (1934), Studies on the helminth fauna of Japan. Pt. 4.
P. auritus auritus has been associated with the gizzard shad. Cestodes or fishes. lap. I. Zoo!., 6, 1-112.
1 Green, J. (1968). The Biology of Estuarine Animals. Sidgwick and Jackson.
Dorosoma cepedianum. However, the larvae have also been London.
found in the mesenteries of the bluegill, Helioperca microchira, ) Thomas, L.J. (1937). On the lire cycle or Contracaecum spicuiigerum.
and of the piscivorous fishes Caenohryllus gluhosus and I.Parasilol" 23,429-431.
4 Chabaud, A. G, (1965). Superfamille des Ascardoidea. In Traite de
Microplerus sppJ. The inference here is that the bluegill, like the
Zoologie, edit. p, P. Grasse. Vol. IV (3), 988-1022, Masson, Paris.
gizzard shad. is the primary source of infestation. which. if eaten
l Kahl, W. (1936). Beitrag zur Kenntnis des Nematoden ContracaeCl4m
by pisdvorous fish. wiIJ release the larvae to enter the cfavatum Rud. Z. Parasitenk" 8,509-520.
mesenteries of the predator where they live as immature ~ Huggins. E, J. (1969). Some parasites of lishes and lishcating birds at Lake
facultative parasites. Ayapel, Columbia, South America,], Parasitol., 55, 539,
South AfricalJ Journal of Science Vol. 73 April 1977 123

High Frequency, High Amplitude


Oscillations in the Amount of O.I.~

'Protein' Extractable from


Cultured Cells
Tncreasing attention is being paid to the oscillatory nature of
ceHular control systems in an effort to explain the cell cycle l - 4
and other dynamic aspects of cell behaviour. 5 - 9 Brodsk y9 has
recently summarised some data on cellular rhythms and, from
studies on amino acid incorporation. polysome and RN A
content and dry weight determinations, has proposed the
existence of rapid nuctuations in the rate of protein synthesis in \j
cells both in viva and in vitro. Although particularly concerned
with periodic variations in enzyme activities and isozyme L-
patterns in relation to differentiation and cancerM , we have 4
undertaken simple spectrophotometric estimations of the protein Time (h)
content of cell extracts. We now wish to report that the level of
extractable protein appears to tluctuate markedly and very Fig. I. Oscillatory variations in the amount of extractable
protein. HaK cells were trypsinised and suspended in their
rapidly under all conditions in all cultured cells examined so far.
filLered medium. After 45-min equilibration at 36.5°C. I ml
Methods aliquots were removed at 5-min intervals and the cells
Human embryonic lung fibroblasts (passage 18). BHK and collected and washed on a filter membrane. After two freeze/
HaK I() hamster cell lines and cells derived from a human thaw cycles the cells were extracted with I ml tris/Hel buffer.
shaken on a vibromix and. after centrifugation. the protein
oesophageal carcinoma 11 were grown in I-litre roller culture
levels were determined from the 280 and 260 nm absorptions
bottles using MEM with Eagles salts (GibCo) supplemented with (details in text). The full curve shows the variations in the values
10 mM HEPES (British Drug Houses) buffer, antibiotics so obtained. while the broken curve gives the changes in the
(200 units/ml penicillin. 100 units/mt streptomycin) and 10% amplitude of the oscillation during the experiment. The latter
foetal calf serum (Flow Laboratories, virus and mycoplasma was determined by joining adjacent peaks (or troughs) of the
screened, heat denatured at 56°C for 30 min). Vervet monkey oscillation and measuring the vertical separation of the
kidney primary cells were supplied by the Vaccine Department intermediate trough (or peak) from this line.
of this Institute as confluent monolayers in a medium consisting
determined. The optical absorbancies (A) of the samples were
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2010).

of OS!'o lactalbumin in Earles salts buffered with bicarbonate.


All cultures were grown to contluency (the HaK cells continued measured at 260 and 280 nm. using tris/HCI butTer as reference.
to proliferate slowly under these conditions) and supplied with and the protein level was estimated from the formula: 12
fresh medium (in some instances without serum when this was protein (mg/ml) - 1.55 X A 2ftU - 0.76 X A260
added during the course of the experiment) approximately 18 h The corresponding enzyme activity and isozyme pattern chooges
prior to preparing the cells for the actual experiment. In order to will not be discussed here but some data have already been
minimise the disturbances to which the cells were subjected, the published..' The effects on the rhythm of 2-phenyl-
medium was filtered through a 0.22.um Sartorius mem- ethanol. EDT A (British Drug Houses) and also dibutyryl
brane to remove debris and free cells. and re-used to cyclic AMP (Boehringer) have so far been examined.
suspend the trypsinised (0.25%, DifCo 1:250, in phosphate
buffered saline. pH 7.8. 0.1 % glucose) cells in a 200 ml BeliCo
suspension vessel fitted with a teflon-coated magnet on a glass
and tetlon shaft. Manipulations were carried out at room
temperature under sterile conditions. After some 45-60 min
equilibration at the experimental temperature (36.5° or 4°C),
sampling from the stirred vessel was begun at regular intervals
(10 min in earlier experiments, latterly at 5 or 2 min). One-m!
aliquots were removed. the cells were collected under suction
and then quickly washed with warm saline (2 ml) on 25 mm "

Sartorius or Millipore filter membranes (0.45 or 1.2 Jlm). The


membranes were then rapidly transferred to liquid nitrogen
(overall sampling time 20-25 s) and stored at this temperature
until required. The cells were then thawed to room temperature
and subjected to a second freeze/thaw cycle. Tris/HCl buffer
(1 ml. 50 mM. pH 7.4) was added and the cells extracted by
•,
"I
shaking on a vibrating mixer for 15 s. Where enzyme activities o 4
were also determined. the extract was frozen and thawed again Time(h)
immediately prior to carrying out the assays. Samples were
Fig. 2. Variations in the mean level of the oscillation in the
generally centrifuged on a MSE angle head bench centrifuge to
amount of extractable protein. The broken curves have been
remove filter and cell membrane debris, although decantation
obtained from the oscillation shown in Fig. I (full curve)
after a short period seemed adequate in some instances. The by joining adjacent peaks and adjacent troughs of the
oscillatory behaviour was observed irrespective of whether or rhythm. The full curve shown here, which indicates the
not these last two steps were carried out but it is possible they variation in the mean level of the oscillation. has been
could slightly influence the observed behaviour. Some samples obtained by taking the midpoints of the vertical distance
were slightly opalescent and thus required treatment so that we between the peaks (or troughs) and the line joining adjacent
prefer to treat all samples where the protein level is to be troughs (or peaks).
124 South African Journal of Science Vol. 73 April 1977

Results some 80-90% at the concentration used (our own unpublished


Figure 1 shows the temporal variations in the level of data). and (iv) to cause the mean level of the lactate
extractable protein in HaK cells determined in the above dehydrogenase activity oscillation in HaK cells (in serum free
manner. A high frequency. high amplitude oscillation is evident medium) to fall by some 75% in 3 h. We are thus of the same
although it is clear that the pattern of behaviour is not constant; opinion as Brodsky'), that the various oscillations are intrinsic
the amplitude, frequency and mean level all changed during the and not induced, although the modulating rhythms may be.
course of the experiment. The dotted curve in this diagram However, we are doubtful that this rapid rhythm involves
shows the apparent changes in the amplitude and it is conduded a periodic change in the rate of synthesis. 9
from this and other comparable data that this parameter can We have also found that the activities and effective isozyme
also vary in a periodic manner. In other words, the observed patterns of several glycolytic enzymess, and also glucose-6-
oscillation is amplitude modulated in a rhythmic way. The phosphate dehydrogenase, vary in an oscillatory manner in all
frequency of the oscillation also appears to vary periodically cells so far examined. These periodicities exhibit the same set of
(rhythmic frequency modulation), at least during the early characteristics as that discussed here, namely frequency,
stages, but settles to a relatively constant value of about 12- 15 amplitude and mean level modulation, often overtly periodic,
min later in the experiment. The subject of frequency modulation and the same resistance to a wide range of agents. 6 Our
has been raised before in relation to the enzyme changes.~ It is preliminary analysis of the data indicates that the periodicity
obvious from Fig. t that the mean level of the oscillation altered discussed here may contribute toward the enzyme oscillations
markedly after some two hours; similar results were obtained in but cannot be entirely responsible for them. *
a duplicate experiment. However, more detailed analysis of the If the oscillation in the level of extractable protein is a universal
data (Fig. 2) indicates that the mean level also fluctuated in a phenomenon. then caution must be observed in the
periodic manner, with a period of about 80 min. Furthermore, it interpretation of at least some specific activity data since
seems feasible for the more marked change at two hours to result experimental ~esults could reflect changes in the total protein
from an even slower variation with a period of the order of content of the extract rather than in the level or activities of the
several hours. n enzymes studied.
These results are typical of those obtained with all the cells This work was supported by a grant from the National
investigated. However, the patterns of modulation exhibited by a Cancer Association of South Africa.
particular cell type apparently reflect both the nature of the cell C. W. A. TSILIMIGRAS
involved and the circumstances. Thus a given modulating signal D. A. GILBERT
mayor may not be overtly periodic but, in general. the Nationalll1stitutefor Virology,
parameters are not constant during the course of an experiment. Private Bag X4,
Sandril1gham 2131,
Discussion South Africa.
Although the method of measurement used is a recommended
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2010).

procedure 12 , it is not yet clear if the observed oscillation is Received December 1.5. 1976.
entirely due to protein level changes. Further work is in progress
to clarify this point. In view of the similarity of these data to the Goodwin, B. C. (1966). An entrainment model for timed enl.yme synthesis
results reported by Brodsky9 on, for example, the rate of amino in bacteria. Nature, 209,479.
acid incorporation, it Seems reasonable to propose that the SeJ'kov. E. E. (1970). Two alternative, self oscillating stationary states
in thiol metabolism- two alternative types of celJ division: normal and
oscillation reflects changes in the amount of extractable protein
malignant ones. Biophysika, 15, 1065.
with time. The mechanism remains obscure. Gilbert, D. A. (1974). The nature of the cell cycle and the control' of
Caution should be exercised in the interpretation of results cel! proliferation. BioSystems, 5, 197.
obtained by discontinuous observation of a periodic Kauffman. S. and Wille. J. J. (1967). The mitotic oscillator in Physarum
phenomenon (especially when its existence has not been poJycephalum. J. theoret. BioI., 55. 47.
, Gilbert, D. A. (1974). The temporal response of the dynamic eel! to
recognised). For example. the apparent frequency can be disturbances and its possible relationship to differentiation and cancer.
dependent on the sampling frequency, giving rise to the problem S. A/r. J. Sci., 70,234.
of aiiasing. 5 For this reason we cannot yet indicate a true period ~ Goodwin, B. C. and Cohcn. M. M. (1969). A phase shift model for

for the oscillation; the involvement of a multiplicity of rhythms spatial and temporal organisation of developing systems. J. theoret.
Bio/., 25,49.
or simple modulation of the oscillation permits one only to give a
Wagner. E., Deill.er. G. F., Fischer, S.. Frosh, S. and Kempf. O. (1975).
possible range for the oscillation frequency. Ideally sampling Endogenous oscillations in pathways of energy transduction as
should be performed more often than we have done here since a related to circadian rhythmicity and photoperiodic control. BioS),stems,
very high frequency component may be present; we note that 7,68.
Tepper et al.l'* have reported that haemoglobin synthesis in Durham. A. C. H. and Ridgway, E. B. (1976). Control of chemotaxis
in physarum po!ycephalum. J. Cell Bio!., 69,218.
reticulocytes occurs in pulses with an interval of some 50 s. In 9 Brodsky, W. Y. (J975). Protein synthesis rhythm. J. theorel. Bio(..
Fig, 2 we have attempted to detect the presence of slower 55, 167.
rhythms by graphical analysis. Elsewhere~ we have referred to 10 Spence, 1. M. (I 963). A. self replicating line from hamster (adult)
an experimental method whereby samples obtained at high k.idney (HaK). S. Afr. J. med. Sci., 28,8 J.
II Bey, E.. Alexander, J., Whitcutt, J. M., Hunt. J. A. and Gear J. H.S.
frequency are combined in groups (filter method). Preliminary
(1976). Carcinoma of the esophagus in Africans. Establishment of
studies of this kind appear to confirm the existence of slower a continuously growing celJ line from a tumour specimen. In Vitro,
rhythms in the amount of extractable protein. 12,107.
The high rrequency oscillation continued in both monkey ,I Layne. E. (1957). Spectrophotometric and turbidometric methods for
kidney ceJJs and HaK cultures at 4°C for at least 5----6 h, the measuring proteins. In Methods in Enzyl7loloKJ', edit. S. P. Colowick and
N. O. Kaplan. Vol. 3, p. 448. Academic Press, New York.
period of observation. At this stage we cannot comment on the
effect of temperature on the frequency other than to say that
* Similar fluctuations have also been detected in the lymphoma eell
there is no gross change. No agent has been found which stops line Raji, which proliferate in suspension. This suggests that the
the high frequency oscillation within the period of observation. existence of the rhythms in other cells is not simply the result of
Even phenylethyl alcohol failed to suppress the protein disturbances arising from trypsinisation. Finally. preliminary studies
oscillation. although it has been reported (i) to dissociate the involving a direct determination of protein in cell extracts by means of
polysomes of rat hepatoma cells u , (ii) to have a cyclic AMP- the Lowry method have yielded an almost identicaJ pattcrn of
like effect 16. (iii) to inhibit the respiration of all these cells by behaviour.
South African Journal of Science Vol. 73 April 1977 12.1
1\ KleveCl. R. R. (969). Tcmporal order in mammalian cells. The periodic 1< Plagernann. P. G. W.. (1968). Phenylethylalcohol: reversible inhibition of
synthesis of lactate dehydrogenase in the cell cycle. J. Cell Bio/., synthesis of macromolecules and disaggregation of polysomes in rat
43.207. hepatoma cells. Biochim. Riophys. Acta. 155,202.
II Tepper. T. Hommes. F. A .. Thurkow. I. and Nijhof. W. (1969). I. Wright, J. A .. Ceri. H .. and Lewis. W. H. (1973). Similar growth of
Oscillations in the rate of haemoglobin synthesis in rabbit reticulocytes. Chinese hamster cells in the presence of phenylethylalcohol or
FFRS Lett., 2. 217. dibutyryl cyclic AMP, NaLUre New Riol., 244.84.

Book Reviews
Enzyme Kinetics A feature of the book which should Atmospheric Physics
Principles of Enzyme Kinetics. By Athel prove valuable is the treatment of complex Atmospheres of the Earth and the Planets.
Cornish-Bowden. Pp.206. (Butterworths. reactions and how to derive steady-state Edited by B. M. McCormac. Pp. 454.
London; 1976.) R24.65. rate equations for them. In addition to his Proceedings of the Summer Advanced
The need for a concise monograph description of how the King/Altman Study Institute. University of Liege,
which treats clearly and rigorously the method is used, Dr Cornish-Bowden Belgium, July 29-August 9, 1974. (D.
fundamentals of enzyme kinetics at an clarifies the principles on which the Reidel, Dordrecht, Holland; 1976.)
intermediate level has been successfully method rests. hitherto not attempted 865.00.
met in this readable book by Dr Comish- outside the original (difficult) paper. In This is another in the series of
Bowden. The author is selective in his this section too are discussed some of the conferences on atmospheric subjects for
choice of material and is at pains to stress practical difficulties in applying the which Billy McCormac is well known.
the importance of understanding the King!Altman method, which may reach There are 32 papers, some of which are
principles of the subject. Limited aspects overwhelming proportions for more didactic reviews. while others are
are covered thoroughly at the expense of a complex mechanisms. Their' simplification straightforward reJXJfts of group research
more encyclopaedic approach in the belief is achieved by the use of newer rules projects. Thus the book has the textbook-
that particular solutions may become derived by the application of kinetics of the cum-journal aspect familiar from its
more complex but not more difficult once theory of flow graphs developed originally predecessors. The papers fall under six
the basics are mastered. Detailed for the analysis of electronic networks. headings: physical processes (7); structure
examples of specific enzymes are Other chapters COVer fast reaction and composition of the neutral and
deliberately omitted so as to emphasize kinetics, the kinetics of control ionized atmosphere (2); laboratory
the general principles involved. Of mechanisms and an analysis of progress measurements of relevant rate
necessity. the subject requires a curves which the author pleads to have coefficients.(4); atmospheres of other
mathematical treatment but clarity is the rehabilitated as a useful enzymological planets (8). Most useful is an introductory
Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2010).

keynote and, as the writer points out in his tool from the neglect into which they section which contains the summaries by
preface, kinetics is not for those who are descended some sixty years ago with the the chairmen of the sessions and an
frightened by the simple algebra and introduction of the Michaelis/Menton authoritative paper by H. 1. Schiff on
calculus required for its understanding. approach. A final chapter summarises neutral chemistry in the earth's
The first chapter is a resume of some of the more useful aspects of atmosphere.
chemical kinetics, followed in successive statIstics for the enzyme kineticist. The conference was in line with recent
chapters by a discussion of the essential Although the application of statistics to thinking in dealing with the whole
characteristics of steady-state enzyme enzyme kinetics may warrant a more atmosphere rather than only a part of it
kinetics. Although these follow the pattern complete treatment, the selection of such as the ionosphere or thermosphere.
as generally taught in numerous material given serves as a valuable Much more attention was paid to overall
biochemical courses, a number of introduction to the subject, particularly on energy sources and sinks than in the past.
viewpoints give a fresh touch to the straight-line fitting. Throughout the book The contributors include such familiar
account. This reviewer was interested. for the reader's attention is constantly names as J .C. G. Walker. D. M. Hunten,
example, in the treatment of V/K m as an directed towards the limitations of the H. Kohl, U. von Zahn, E. E. Ferguson, G.
important parameter in its own right (at methods used and the reliability which Kockarts, J. S. Nisbet, W. Swider, P.
least as imJXJrtant as that of Km) and the may be placed on the assumptions Stubbe, G. G. Shepherd. T. M. Donahue,
discussion of the vexing problem of the inherent in the mathematical treatment. M. H. Rees, D. F. Strobel and M. B.
best linear plot of the Michaelis/Menton The dangers of using concepts without a McElroy.
relationship. Possibly. however, the clear understanding of such limitations is It is a pity that the volume only reached
author is shouting into the wind when stressed and, while principally a this reviewer in December 1976, for this
trying to discourage the use of the theoretical treatment, the coverage is well means inevitably that many of the more
Lineweaver-Burk plot which he shows to geared towards solving the problems recent data are already Q.Utmoded. The
be the least satisfactory of a number of which arise in the handling of real data. chief permanent interest of the volume
possibilities. I enjoyed this book and feel sure that it rests in the reviews on such subjects as
Enzyme inhibition is considered in will receive an appreciative audience. not atmospheric and ionospheric modelling,
some detail and it is interesting to note the only among the first-year research and the atmospheres of Mars, Venus and
virtual dismissal of non-competitive students at which it is aimed, but also Jupiter. Most of the material is available
inhibition as an important type of enzyme other investigators who use kinetic elsewhere in the literature and some needs
phenomenon. Likewise, the effect of methods. no matter how limited, and wish updating.
temperature on enzyme-catalysed to have a readable and authoritative The book is a useful reference for
reactions. though its importance is account of the principles involved in this workers in the appropriate fields, but even
aCknowledged, is given short shrift. "So many-sided subject. at the present inflated rates the price is
Few of the many temperature studies G. E. Trout very high. J. A. Gledhill
carried out on enzymes are of any value." Department of Physiology and Medical Department of Physics.
points out Dr Cornish-Bowden. Biochemistry. UnNersity of Cape Town. Rhodes Uni~'ersity. Grahamstown.

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