42 Nature Vol.

259 January I & 8 1976

19 Crawshaw, L. 1., Comp. Biochem. Physiol., SlA, 11-14 (1975). and made progress back towards the pouch. The joey was
20 Neill, W. H., and Magnuson,J.J., Trans. Am. Fish. Soc.,lOJ, 663-710 (1974).
21 Beitinger, T. L., Fish. Bull. Mass., 11., 1087-1093 (1974). not handled during this experiment.
22 Neill, W. H., Magnuson, J. J., and Chipman, G. D . , Science, 176, 1443-144'
(1972). These observations suggested that the newborn quokka
has a righting reflex; we found that it could also turn over
when placed on its back. But older pouch young, up ·to
the age of 65-70 d, no longer retain this ability, and
Gravity as the sole histological sections through the head of a newborn joey
support the conclusion that the vestibular apparatus is
navigational aid to the newborn quokka undifferentiated'·". It seems to us that this transient right-
ALTHOUGH there is general agreement on the sequence of ing reflex could be mediated by muscle stretch receptors,
events at birth for several marsupial species', it is still not particularly in the neck.
known how the newborn joey navigates on its journey These experiments confirm the negative geotropism of
from the mother's cloaca to her pouch. Frith and Calaby' the newborn quokka and reveal no other mechanism
have postulated that, at birth, the red kangaroo (Megaleia directing its movement towards the pouch. In the second
rufa) ". . . has large nostrils and presumably a well experiment, in particular, the joey passed within I em of
developed sense of smell which probably plays a major the pouch opening without hesitating in its upward climb
part in its location of the pouch". On the other hand, and it is clear that chemotaxis can be discounted in this
Hartman' has suggested that the newborn opossum case.
(Didelphis virginiana) was "negatively geotropic" though We thank Professor A. R. Main for helpful discussions.
he has now modified this view considerably•, as histological ]. R. CANNON
studies have revealed that the vestibular apparatus of the Department of Organic Chemistry
newborn animal is undifferentiated and therefore non- H. R. BAKKER
functional'·•. To investigate whether one of these s. D. BRADSHAW
mechanisms' ·• could account for the behaviour of the new- Department of Zoology,
born quokka (Setonix brachyurus) we have observed par- University of Western Australia,
turition in this macropodid marsupial. Nedlands, Western Australia
Female quokkas mate soon after giving birth and such I. R. McDoNALD
animals often develop a blastocyst in the state of diapause Department of Physiology,
in the uterus. Removal of the pouch young causes this Monash University, Victoria, Australia
blastocyst to resume development so that a birth occurs Received September 9; accepted November 13, 1975.
about 26 days Jater 7 • In March 1974 and 1975 pouch 1 Tyndale-Biscoe, H., Life of Marsupials (Edward Arnold, London, 1973).
young were removed from a total of 14 quokkas obtained 2 Frith, H. J., and Calaby, J. H., Kangaroos (Cheshire, Melbourne, 1969).
from a wild population on Rottnest Island, Western 3 Hartman, C. G., A nat. Rec., 19, 251 (1920).
• Hartman, C. G., Possums, cb. 13 (University of Texas Press, Austin, 1952).
Australia. The animals were transferred to a controlled s Larsen, 0., McCrady, E., Jr, and Zimmermann, A. A., J. comp. Neurol. 63, 95
(193').
environment 22 d later, then observed continuously. In all, 6 Langworthy, 0. R., Am. J. Physiol., 74, I (1925); J. comp. Neuro/., 46,201 (1928).
7 Shield, J . W., and Woolley, P., Nature, 188, 163 (1960}.
seven animals gave birth; four births were seen and one 8 Tyndale-Biscoe, C. H., thesis, Univ. Western Australia (1961).
was recorded in detail on videotape.
Our observations of these births confirm those of
Tyndale-Biscoe•. In spite of intensive antepartum groom-
ing of the pouch and cloacal region, there is no evidence
of a path made by the mother to facilitate the passage of Impaired acquisition of a passive avoidance
the joey from the cloaca to the pouch. The mother's posi- response after lesions induced
tion during birth was such that the abdominal wall was in the locus coeruleus by 6-0H-dopamine
almost vertical and the lower lip of the pouch pouted
outwards. Observation of the actual entry of the joey into CRow' and Kety' have advanced the hypothesis that the
the pouch was difficult because of the stance of the mother fine network of noradrenaline-containing nerve terminals
but on one occasion the joey seemed merely to tumble in which innervates the cerebral cortex' is a necessary com-
over the lip. ponent of the mechanism of learning. In particular, it is
After a birth in 1974 the joey was retrieved as soon as it suggested that these neurones deliver a "results of action"'
had reached the pouch. The mother was then held ver- signal which registers the successful outcome of a particular
tically, head up, and the joey was placed on the upper behavioural sequence. The noradrenaline released in the
(anterior) lip of the pouch. During the next 3 min it cortex is believed to interact with a transient synaptic
climbed about 3 em upwards away from the pouch. The change resulting from recent neural activity (the 'short-term
mother was then turned upside down. The joey faltered trace'), to initiate the long term synaptic changes presumed
and eventually turned completely around so that its head to underlie learning. Various authors have suggested the
was again pointed upwards, towards the pouch. It then existence of such a "confirming reaction"', "results of
became entangled in the mother's fur and made no pro- action'", or "now print"" mechanism. The diffuse distribu-
gress. When the mother was returned to the head up tion of noradrenergic .terminals in the cerebral cortex would
position the joey reversed its direction again and pro- allow this system to deliver a generalised message of the
gressed away from the pouch. It was then removed from type required. Ungerstedt' has demonstrated histochemi-
the mother's chest and replaced near the cloaca, whence cally that the cortical noradrenergic terminals originate
it succeeded in making a second .iourney to the pouch. from cell bodies of the nucleus locus coeruleus in the dorsal
On the occasion of the videotaped birth in 1975 the pontine tegmentum, and this has been confirmed in bio-
mother was removed from her cage and held almost ver- chemical studies•·•. The concept of the coerulo-cortical
tical, with her right side slightly lower than her left, while noradrenergic system as a reinforcement pathway is sup-
the newborn joey was sti·ll making its initial journey to the ported by the observation that rats with electrodes in the
pouch. This led to its direction changing slightly so that region of the locus coeruleus can be trained to self
it passed by the pouch; it did not falter but continued to stimulate"·" and this behaviour is accompanied by in-
climb to the mother's chest, leaving a distinct path, about creased noradrenaline turnover in the ipsilateral cortex 12 •
I em deep, in the fur. The mother was then turned to the The prediction that bilateral lesions of the locus coeruleus
head down position and, as before, the joey slowly turned would impair learning capacity'·'·" has been tested in a
around, this time without becoming entangled in the fur, runway' and a two-choice discrimination learning test>' .
© 1976 Nature Publishing Group