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Thalamic projections to the somatosensory cortex of the echidna, Tachyglossus aculeatus.

Evoked potential studies (Lende, '64) suggest that echidnas have a single, topographically organized somatosensory area (SMI) that spans a mediolaterally oriented sulcus called sulcus alpha. A motor area (MI) is situated on the prealpha gyrus. This study examines the cytoarchitecture and thalamic afferents of SMI in the echidna, Tachyglossus aculeatus. SMI contains two cytoarchitectonic fields. A caudal field extends across the postalpha gyrus and onto the floor of sulcus alpha. It has a well-developed layer 4 and a relatively small number of medium-sized pyramidal cells in layer 5. The rostral field extends from the floor of sulcus alpha onto its rostral bank. It also has a well-developed layer 4 but has a large number of large pyramidal cells in layer 5. Layer 4 thins as it is followed onto the crown of the prealpha gyrus. The remainder of this gyrus contains a single cytoarchitectonic field with a thin layer 4 and a layer 5 heavily populated with larger pyramidal cells. This field corresponds to the physiologically defined motor area MI. Thalamic afferents to SMI were examined by placing pressure injections of horseradish peroxidase into the two cytoarchitectonic fields. An injection that involved both fields retrogradely labeled neurons throughout the ventral posterior nucleus of the thalamus. An injection restricted to the caudal field labeled a band of neurons that extends rostrocaudally throughout the ventral part of the ventral posterior nucleus. An injection restricted to the rostral field labeled a band of neurons situated dorsally in the ventral posterior nucleus. No other thalamic groups contained labeled neurons comparable to the labeling seen in the intralaminar or posterior nuclei following a horseradish peroxidase injection into SI of marsupial or placental mammals. These results indicate that SMI in Tachyglossus contains two cytoarchitectonic fields that resemble areas 3a and 3b in some placental mammals, suggesting that the constellation of cytoarchitectonic fields corresponding to areas 4, 3a, and 3b is a basic mammalian character which has been modified in marsupial and many placental mammals.

Afferent Pathways↗

Morphology and distribution of neurons in the retinal ganglion cell layer of the adult tammar wallaby--Macropus eugenii.

The morphology of the ganglion cell layer of the adult tammar wallaby has been examined from Nissl-stained retinal flatmounts. From this material, neurons have been classed as ganglion cells or displaced amacrine cells according to the disposition of Nissl substance. A further subdivision of ganglion cells into a separate group of alphalike cells was assisted by determining the range of soma sizes in neurofibrillar-stained flatmounts, a method which, in the cat, has revealed the presence of alpha cells. Isodensity contour maps prepared from the Nissl-stained flatmounts show a well-developed visual streak and an area centralis in the total neuronal population. A similar pattern was also found in the ganglion cells, thus confirming Tancred's (J. Comp. Neurol. 196:585-603, '81) finding, and, as well, in the alphalike ganglion cells and the displaced amacrine cells. The relative proportions of ganglion cells to displaced amacrines (GC:DA) were evaluated from isodensity profiles drawn along and vertical to the visual streak for the two cell types and also from maps showing the variation in the GC:DA ratio throughout the retina. A comparison with results published for other species shows that the visual streak development in the tammar wallaby is consistent with the expectations of the "terrain" theory and that, in its relative proportion of displaced amacrines, the tammar closely resembles the rabbit but contrasts sharply with the cat, which has half as many ganglion cells and three times as many displaced amacrines as the other two species.

Animals↗

Mammalian motoneuron cell death: development of the lateral motor column of a wallaby (Macropus eugenii).

We have investigated the development of the lumbar lateral motor column of the tammar as a model of mammalian motoneuron cell death that is accessible to experimental manipulation. The tammar is an Australian marsupial, belonging to the subfamily of wallabies and kangaroos. After a gestational period of 26-28 days, the pup crawls to its mother's pouch using its forelimbs. The major morphometric events that shape the formation of the hindlimb occurred between 21 days gestation and birth. At birth the premuscle masses had divided and motor nerves had begun to penetrate the muscles of the thigh and shank. The period of motoneuron cell death was biphasic and occurred entirely postnatally. During phase I, between birth and 40 days, 59% of motoneurons were lost. Cell numbers then stabilised before falling a further 24%, to give an overall loss of 70%. Most of phase II cell loss occurred between 90 and 150 days. The possibility that a second period of motoneuron cell death may be a common feature of mammals is discussed.

Animals↗

Retinotopic organization in the dorsal lateral geniculate nucleus of the tammar wallaby (Macropus eugenii).

Electrophysiological recordings were made from 187 single cells in the tammar wallaby (Macropus eugenii) dorsal lateral geniculate nucleus (LGNd). The results show that it is topographically organized such that the superior visual field is represented dorsally, the inferior field is represented ventrally, the nasal visual field is represented caudally, and the temporal visual field is represented rostrally. The visual field of one eye ranges from -30 degrees nasal to +179 degrees temporal in azimuth and +73 degrees superior to -49 degrees inferior in elevation. Ganglion cells that had receptive field positions between -9 degrees and +179 degrees projected to the contralateral LGNd while the ganglion cells that projected to the ipsilateral LGNd had visual fields from 0 to +30 degrees. The binocular visual field extends 60 degrees in azimuth. This representation in the LGNd is expanded relative to the monocular representation. There is also an increased representation of the horizon in the temporal field corresponding to the visual streak of retinal ganglion cells. The binocular visual field is located where contralateral and ipsilateral laminae are shown to interdigitate by proline autoradiography. There are nine eye-specific laminae in the LGNd. Four receive afferents from the contralateral eye and five receive afferents from the ipsilateral eye. The lines of isoelevation are perpendicular to the coronal plane of section while the lines of isoazimuth are nearly parallel to the coronal plane. The lines of projection representing one visual direction are inferred to be perpendicular to the tangent of curvature of the laminae as in the LGNd of other mammals. The majority of cells (85%) recorded had on- or off-centre responses. On- and off-centre responses were not apparently segregated in the LGNd but segregation may not have been revealed by the single-unit recording technique.

Animals↗

Cell death in the developing retinal ganglion cell layer of the wallaby Setonix brachyurus.

The distribution and number of dying cells in the developing retinal ganglion cell layer of the wallaby Setonix brachyurus were assessed by using cresyl violet stained tissue. The density of dying cells has been expressed per 100 live cells for the entire retinal surface, data being presented as a grid of 500 micron squares. For statistical analysis, retinae were divided into 8 regions; dorsal, ventral, nasal, and temporal quadrants, each further divided into center and periphery. This method allowed comparison of the extent of cell death at different retinal locations as the high density area centralis of live cells developed temporal to the optic disk from 60 days onward. Between 30 and 70 days, dying cells were seen across the entire retina; beyond 100 days very few were seen. Initially, there was a significantly higher incidence of dying cells in the central retina compared to the periphery, whereas from 50 days this situation was reversed. Analysis of the central retina before and during area centralis formation consistently indicated a significantly lower number of dying cells per 100 live cells in temporal compared to other retinal quadrants. This differential pattern suggests that cell death lowers live cell densities less in the emerging area centralis than elsewhere, and therefore must play a part in establishing live cell density gradients. However, we cannot exclude the possibility that other factors are also instrumental. Indeed, factors such as areal growth (Beazley et al., in press) presumably operate at later stages since live cell density gradients continue to be accentuated even after cell death is complete. Numbers of dying cells peaked by 50 days, reaching approximately 1% of the live cell population. At this stage, counts were also maximal for live cells with values up to 30% above the adult range.

Aging↗

Mammalian motoneuron development: effect of peripheral deprivation on motoneuron numbers in a marsupial.

In nonmammalian vertebrates, the survival of developing motoneurons is dependent on their contacting appropriate target cells. It is generally accepted that developing mammalian motoneurons have a similar dependency on their target, but as yet there is little experimental evidence to support this contention. This is mainly because of the difficulty of experimenting on eutherian embryos. We have, therefore, been studying neuronal development in the tammar (an Australian marsupial) as its nervous system is immature at birth. Radical or partial removal of hindlimb buds from newborn tammars resulted in an increased motoneuron cell death. The motoneurons which survived in the operated tammars did so by innervating muscle remnants. In the instances where a group of muscles was totally removed, the corresponding motonuclei appeared to be totally lost. This study supports the hypothesis that mammalian motoneurons must contact their appropriate muscle in order to survive through the period of natural neuronal cell death.

Age Factors↗

Retinal projections to the superior colliculus and dorsal lateral geniculate nucleus in the tammar wallaby (Macropus eugenii): I. Normal topography.

The topography of retinal projections to the superior colliculus and dorsal lateral geniculate nucleus of a wallaby, the tammar (Macropus eugenii), was investigated by an anatomical method. Small laser lesions were made in the retinas of experimental animals, and the remaining retinal projections were visualized by means of horseradish-peroxidase histochemistry. The position of each lesion was correlated with the position of the filling defects in the terminal label. The whole of the retina projects to the contralateral superior colliculus. The nasal retina is represented caudally, and the temporal retina rostrally. The ventral retina is represented medially, and the dorsal retina laterally. There is a projection to the ipsilateral superior colliculus, but it is patchy and its topography could not be determined by this method. The retinotopic map in the contralateral dorsal lateral geniculate nucleus has the nasal retina represented rostrally and the temporal retina caudally in the nucleus. The dorsal retina is represented ventrally, and the ventral retina is represented dorsally. It appears that the whole of the retina projects contralaterally, and in addition the temporal retina projects ipsilaterally. The maps of visual space through the two eyes were shown to be in topographic register in the binocular region by making a deposit of HRP in the visual cortex. This resulted in a column of retrogradely labeled cells in the nucleus. This column crossed the laminae, which are innervated by the ipsilateral and contralateral eye at right angles.

Animals↗

Retinal projections to the superior colliculus and dorsal lateral geniculate nucleus in the tammar wallaby (Macropus eugenii): II. Topography after rotation of an eye prior to retinal innervation of the brain.

Retinal projections to visual centers in a marsupial mammal, the tammar wallaby (Macropus eugenii), have been investigated after an eye rotation prior to retinal innervation of the brain. Retinal topography to the superior colliculus and dorsal lateral geniculate nucleus was mapped by using laser lesions of the retina and horseradish peroxidase histochemistry. Despite the change in orientation of optic axon outgrowth from the developing eye after rotation, retinal ganglion cells made orderly connections in the colliculus and geniculate according to their original retinal position within the eye and not their rotated position. Axons must have corrected their pathways at some point between the back of the eye and their targets. The optic chiasm was one such site. Optic axons from the rotated eye took an abnormal course at the caudal end of the chiasm. Growth of optic axons through aberrant pathways in the brain did not preclude specific innervation of targets. When by chance optic axons entered through the oculomotor nerve root they specifically innervated their correct visual centers, albeit in reduced density, and did not innervate inappropriate targets. These results support the idea of specific interactions between growing axons, the pathways they grow along, and their targets.

Animals↗

The accessory optic system of the wallaby, Setonix brachyurus: anatomy in normal animals and after early unilateral eye removal.

We have traced primary visual projections to nuclei of the accessory optic system in the mature wallaby, Setonix brachyurus, the "quokka," following unilateral intraocular injections of horseradish peroxidase. The organization of pathways and nuclei is similar to that of other marsupials and to that of eutherian mammals. The dorsal, lateral and medial terminal nuclei receive bilateral input, though nuclei ipsilateral to the injected eye are weakly labelled in comparison with their contralateral counterparts. We also report on the accessory optic system in animals which were unilaterally enucleated neonatally or at postnatal day 35. At maturity in enucleated animals, ipsilateral projections to all nuclei of the accessory optic system are more densely labelled than normal. This exuberance is more pronounced in neonatally enucleated animals than in those enucleated at the later stage.

Animals↗

Auditory forebrain organization of an Australian marsupial, the northern native cat (Dasyurus hallucatus).

Structures and connections of auditory forebrain regions of the Northern native cat, a member of one of the most primitive families among Australian marsupials, have been examined anatomically by using anterograde and retrograde tracing techniques with wheat germ agglutinin-conjugated horseradish peroxidase (WGA-HRP) after defining the acoustically responsive neocortical area physiologically. The structure of the medial geniculate body (MG) was similar to that described in other species. The results obtained from a case with a WGA-HRP injection into the MG showed that the MG strongly projects to the lateral amygdaloid nucleus (LAmy) and the putamen as well as the auditory neocortex (ACx). Results obtained from other cases with WGA-HRP injections into the physiologically defined ACx show also that the ACx is connected not only with the ipsilateral MG and the contralateral ACx but also with the LAmy both bilaterally and reciprocally. The regions within the LAmy in which the MG-LAmy projection fibers terminate largely overlap with those in which the ACx-LAmy fibers terminate and the LAmy-ACx pathway originates. The connectional relationships revealed in the present study--that the LAmy receives auditory information from the MG and reciprocates auditory information with the ACx bilaterally--strongly suggest that, in some primitive mammals with small neocortical areas, a specific portion of noncortical telencephalon functions as an auditory center and occupies a relatively large volume of space in the forebrain. It is possible that the auditory sector of noncortical telencephalon in some primitive mammals such as the American Didelphidae and the Australian Dasyuridae is homologous with part of the auditory sector of the dorsal ventricular ridge (DVR) in reptiles and birds and also may have functions shared with the auditory primary and association neocortex in advanced mammals such as the domestic cat and the monkey (Kudo et al., '86a).

Animals↗

Effects of very early monocular and binocular enucleation on primary visual centers in the tammar wallaby (Macropus eugenii).

The role of retinal afferents and their binocular interactions in the development of mammalian primary visual centers has been studied in the marsupial wallaby. Monocular and binocular enucleation was performed prior to any retinal innervation of the visual centers. After monocular enucleation retinal projections were traced by horseradish peroxidase histochemistry and compared with those in normal animals and those during development. The topography of retinal projections to the superior colliculus and the dorsal lateral geniculate nucleus after monocular enucleation was determined by making retinal lesions and tracing the remaining projections with horseradish peroxidase. The position and nature of the filling defects in terminal label were compared with controls with similarly placed lesions. The superior colliculus and dorsal lateral geniculate nucleus ipsilateral to the remaining eye were shrunken. Projections to the ipsilateral superior colliculus, ipsilateral accessory optic nuclei, and ipsilateral suprachiasmatic nucleus, although enlarged, never approached the density contralaterally, as was also the case during normal development. The expanded projection in the ipsilateral superior colliculus came primarily from temporal and ventral retina. In the dorsal lateral geniculate nucleus, terminal bands and cellular laminae, although not identical to normal, did develop. During normal development overlap of afferents from the two eyes occurs in the binocular region. The decrease in volume of the nucleus ipsilateral to the remaining eye after monocular enucleation suggests that the monocular region disappears in the absence of appropriate input and the binocular region survives. Contralaterally there was no decrease in volume, compatible with this idea. The topography of retinal projections supports this interpretation. It was normal contralaterally while ipsilaterally it was appropriate for the normal binocular region. There was an expansion of the projection along the lines of projection in what would normally be binocular regions of the nucleus, where retinal afferents failed to segregate in the absence of binocular competition. After binocular enucleation the alpha and beta segments of the dorsal lateral geniculate nucleus were still recognizable but cell-sparse zones were absent, as was the characteristic orientation of primary dendrites of geniculocortical cells. There are rigid developmental constraints operating on the innervation of territory by retinal afferents from the two eyes, and many features of the mature pattern arise without binocular interactions during development.

Afferent Pathways↗

Cell death in the inner and outer nuclear layers of the developing retina in the wallaby Setonix brachyurus (quokka).

We have examined the number and distribution of dying cells in the developing inner (INL) and outer (ONL) nuclear layers of sectioned quokka retinae (N = 31) from embryonic day (E)24 to postnatal day (P)192. Before birth, dying cells were seen in the optic fissure. Thereafter two major phases of cell death took place in the INL. The first phase was more pronounced within the vitread part with peak numbers of dying cells at P50. By contrast, during the second phase, cell death was more extensive in the sclerad portion; peak numbers of dying cells were recorded at P85 and P100 for the vitread and sclerad parts respectively. At these stages, photoreceptors were seen in the INL suggesting that these ectopic cells contribute to the pool of dying cells. The pattern of cell death broadly followed a central to peripheral sequence in the first phase but, in the second, was seen initially in mid-temporal retina and then became panretinal. Dying cells were seen in the ONL but in smaller numbers than in the INL. There was a peak of cell death at P26 which may represent death of mitotic cells at the ventricular surface. In the quokka, retinal cell genesis takes place in two phases (Harman and Beazley: Neuroscience 28:219-232, '89). The two major phases of cell death described here peak approximately 40 days after episodes of maximal cell genesis. These findings, together with data for the mouse, suggest that a biphasic pattern of cell genesis and cell death may be a feature of eutherian as well as marsupial retinal development.

Aging↗

Early development of retinal ganglion cell dendrites in the marsupial Setonix brachyurus, quokka.

The dendritic morphology of retinal ganglion cells was studied in flat-mounted retinae of the marsupial Setonix brachyurus, quokka. In the adults, horseradish peroxidase (HRP) was applied to the vitread surface of flattened retinae. Wide-, large-, medium-, and small-field classes appeared to correspond to gamma, alpha, delta, and beta cells, respectively, in the cat (Boycott and Wässle, J. Physiol. 249:397-419, 1974). To reveal the early stages of dendritic development, HRP was placed on the optic nerve of isolated eye cups from the day of birth to postnatal day (P) 63 when the area centralis is beginning to form (Dunlop and Beazley, Dev. Brain Res. 23:81-90, 1985). Youngest cells lacked dendrites and had an elongate soma in the cytoblastic layer with an endfoot contacting the ventricular surface. Once in the ganglion cell layer, the soma was rounded and dendrites appeared as short, unbranched processes. Most cells were asymmetric or "polarised" with the axon arising from the side nearest the optic disk and dendrites from the opposite side. Polarity was maintained in cells with longer, branched dendrites. A small proportion of cells exhibited a reversed polarity in which the axon arose from the side nearest the retinal edge and dendrites towards the disk. Cells appeared to acquire an approximately symmetric, adult-like tree by the addition of new primary dendrites between the existing ones and the axon hillock. Wide-, large-, medium-, and small-field cells were evident from P6, P25, P31, and P40, respectively. Spines were observed on dendrites and axons during development but were rare in the adult. Some dendro-axons were seen at all ages examined. The existence of an initial axodendritic polarity in retinal ganglion cells supports the hypothesis that the axon hillock is the determinant of dendritic geometry (Maffei and Perry, Dev. Brain Res. 41:185-194, 1988). Polarity may also contribute to the establishment of "radial orientation" in which the long axis of the elliptical dendritic tree of cells outside the area centralis points towards central retina and the weighted centre is displaced towards the retinal periphery (Leventhal and Schall, J. Comp. Neurol. 220:465-475, 1983).

Animals↗

Development of retinotopy in projections from the eye to the dorsal lateral geniculate nucleus and superior colliculus of the wallaby (Macropus eugenii).

The development of retinotopy in projections from the eye to the dorsal lateral geniculate (dLGN) and superior colliculus (SC) has been studied in the marsupial wallaby. Discrete retinal lesions were made and the remaining retinal projections were traced with horseradish peroxidase in animals at stages ranging from just after optic innervation of the dLGN and SC to the time when the projections are mature. Topographically organised projections could be recognized a few weeks after axons first reached the dLGN and SC with a topographically discrete projection from nasoventral retinal recognized later than from dorsal, dorso-temporal, temporal, and temporoventral retina. Over time there was an increase in precision of the retinotopy as judged by an increase in sharpness of the borders of filling defects in the projection labelled with horseradish peroxidase. Refinement of the projection from temporal retina preceded that from nasal retina in both the dLGN and SC and in the former occurred concomitantly with the segregation of eye-specific terminal bands. Refinement was complete 16 weeks after birth, prior to eye opening at around 20 weeks after birth. Inequalities in retinal representations in both nuclei were present from the time retinotopy could first be detected. This was before the inequalities in retinal ganglion cell distribution, which underly these representations in the adult, were obvious. Retinotopy and inequalities in retinal representation characteristic of the adult are present from a very early stage in the protracted development of visual projections in the wallaby. Refinement may involve death of inappropriately projecting cells, pruning of inappropriately projecting axon arborizations or could be achieved by growth of the retinorecipient neuropil. Temporonasal differences in the time course of refinement may reflect gradients of maturation in the retina.

Animals↗

Neurogenesis in the brain auditory pathway of a marsupial, the northern native cat (Dasyurus hallucatus).

Neurogenesis in the auditory pathway of the marsupial Dasyurus hallucatus was studied. Intraperitoneal injections of tritiated thymidine (20-40 microCi) were made into pouch-young varying from 1 to 56 days pouch-life. Animals were killed as adults and brain sections were prepared for autoradiography and counterstained with a Nissl stain. Neurons in the ventral cochlear nucleus were generated prior to 3 days pouch-life, in the superior olive at 5-7 days, and in the dorsal cochlear nucleus over a prolonged period. Inferior collicular neurogenesis lagged behind that in the medial geniculate, the latter taking place between days 3 and 9 and the former between days 7 and 22. Neurogenesis began in the auditory cortex on day 9 and was completed by about day 42. Thus neurogenesis was complete in the medullary auditory nuclei before that in the midbrain commenced, and in the medial geniculate before that in the auditory cortex commenced. The time course of neurogenesis in the auditory pathway of the native cat was very similar to that in another marsupial, the brushtail possum. For both, neurogenesis occurred earlier than in eutherian mammals of a similar size but was more protracted.

Aging↗

Visual factors in sound localization in mammals.

The ability of mammals to localize sound varies widely among species. During the past decade, evidence has accumulated that this variation cannot be accounted for simply on the basis of the availability of the physical cues for locus. Evidence is presented that a major factor in sound localization is the need to direct the field of best vision to a sound source for further scrutiny. Thus, species with broad fields of best vision (such as visual streaks) require less accurate information regarding the location of a sound source than do species with very narrow fields of best vision (such as foveae). To support this suggestion, data are reported for the width of the field of best vision in the form of retinal ganglion cell isodensity contours for thirteen species of mammals. The possible contribution of other factors including binocular fields, visual acuity, and the degree to which a species is predatory in lifestyle, is also examined.

Animals↗

The morphological development of the inferior colliculus in a marsupial, the Northern quoll (Dasyurus hallucatus).

As a part of a project concerning the development of hearing, some features of the morphological development of the inferior colliculus were studied in a marsupial, the Northern quoll or native cat (Dasyurus hallucatus). Marsupials are of particular interest in developmental studies because much embryonic development occurs outside the uterus, in the pouch. Nissl-stained material was prepared from pouch-young at various ages between 11 and 81 days, and for a number of adults. Four pouch-young were injected with tritiated thymidine and killed later during pouch life. The inferior colliculus is first recognizable in pouch-young aged 23 days, when it is bordered by a cell-sparse ring of tissue. By this time, the labelling patterns following injections of tritiated thymidine made on days 7-9 suggest that migration of cells to the inferior colliculus from the ventricular germinal zone has been largely completed. At 81 days, close to the time when the young move out of the pouch, the adult cytoarchitecture--a central nucleus flanked by dorsal and lateral cortical regions--is clear. Cell areas expand monotonically as a function of age. The period of days 45-50 is associated with a large expansion of cell volume and a concomitant decrease in packing density. It is likely that functional connections are forming during this period, which may herald the onset of hearing in the quoll. Total cell numbers increase to a peak at day 36, fall to a minimum at day 50, and rise again to the adult value. The second increase is likely to be a phase of glial proliferation, in part associated with the onset of myelination. This increase correlates with departure of the young from the pouch.

Aging↗

Hearing, vocalization and the external ear of a marsupial, the northern Quoll, Dasyurus hallucatus.

As part of a continuing study of the development of the marsupial auditory system, auditory brainstem responses (ABR) were recorded and an ABR audiogram was constructed for five female Northern Quolls (Dasyurus hallucatus), which are nocturnal carnivores. The best frequency for hearing lies between 8 and 10 kHz, and at 50 dB SPL there is a range from about 0.5 to 40 kHz. Vocalizations of adult quolls and pouch-young were recorded with a digital audio tape recorder, and the power spectra of representative calls were compared with the ABR audiogram. The common adult vocalizations have most energy at the lower end of the hearing range, whereas frequencies that are dominant in the isolation calls of the pouch-young lie close to the best frequency of hearing. Samples of nocturnal sounds of the habitat of the quoll were also recorded and analyzed. Power spectra have peak energy at frequencies between 2 and 5 kHz, with a smaller contribution above 10 kHz. The spectrum contains relatively little power at the best frequency of hearing. Measurements of the sound pressure level at the external ear canal as a function of stimulus frequency and location in space suggest that the directional amplifying properties of the pinna will operate most effectively on sound frequencies at the upper end of the quoll's hearing range, a region that may be important in prey detection. Comparisons are made with other mammalian nocturnal carnivores and with other marsupials. We speculate that, for nocturnal carnivores, one role of the low-frequency part of the hearing range concerns the recognition of adult conspecifics, the mid-frequency range is important for the detection of pouch-young, and the upper range may be particularly concerned with prey/predator detection.

Animals↗