Tree Shrew Tupaia Research Articles

Cranial circulation of the pen-tailed tree shrewPtilocercus lowii and relationships of Scandentia

The major cranial arteries and veins are described for a 30-mm crown-rump length fetus of the pen-tailed tree shrewPtilocercus lowii, and comparisons are made with cranial vessels reported in the tree shrewTupaia and with the vascular pattern reconstructed for primitive eutherians.Ptilocercus shares a number of derived features of the cranial circulation withTupaia, which, therefore, represent synapomorphies of tree shrews (Tupaiidae, Scandentia). Included are (1) the enclosure of the intratympanic portion of the internal carotid artery in a bony canal that is floored proximally and distally by the entotympanic and by the petrosal in between, (2) the enclosure of the intratympanic portion of the stapedial artery by the petrosal in a canal on the promontorium and within the epitympanic crest beneath the tympanic roof, (3) the absence of an exit for the arteria diploetica magna, (4) an alisphenoid canal, (5) a maxillary artery that passes medial to the mandibular nerve beneath foramen ovale, and (6) a laryngeopharyngeal artery. Some of these derived features, however, are also found in certain other eutherians (e.g., numbers 2, 3, and 6 in Euprimates) and, therefore, may be used in future studies to assess the higher-level affinities of Scandentia.

Immunocytochemical identification of photoreceptor populations in the tree shrew retina

The presence and retinal distributions of short-wavelength-sensitive cones, long-wavelength-sensitive cones, and rods were assessed in the retina of the tree shrew Tupaia belangeri using visual pigment antibodies OS-2, COS-1 and anti-rhodopsin, respectively. Results demonstrated a clear immunocytochemical differentiation of the three photoreceptor types with each showing regional variation in density across the retina.

Interlaminar connections of the superior colliculus in the tree shrew. I. The superficial gray layer.

One of the most persistent problems in the study of the superior colliculus is the relationship between its superficial and deep layers. The superficial tier of layers is considered to be visuosensory in function, whereas the deep tier is multisensory and has premotor functions. This fundamental distinction is the primary basis for the view that a visually triggered shift in the direction of gaze depends on the transfer of information from sensory cells in the superficial tier to premotor cells in the deep tier. The goal of the present experiments was to examine the interlaminar projections of the superficial gray layer in the tree shrew Tupaia belangeri. We used biocytin as the marker for tracing the pathways. The tree shrew was chosen because its large and distinctly laminated superior colliculus facilitates the task of examining connections between the layers. Biocytin was used because of its sensitivity and because it allowed us to place very small injections restricted entirely to the superficial gray layer. The results demonstrated that a prominent pathway originates in the superficial gray layer and terminates in stratum opticum. In comparison, the projection from the superficial gray layer to the layers beneath stratum opticum is extremely sparse. The pathway from the superficial gray layer to stratum opticum has a columnar distribution, extending about 100 microns rostrally and caudally from the center of the injection site. There were no signs of more remote intracollicular connections, nor of patches or bands of terminals. The biocytin injection sites also labeled pathways to nuclei as distant from the superior colliculus as the diencephalon, including the dorsal and ventral lateral geniculate bodies, and the pulvinar. The results suggest that stratum opticum may serve as a link between the superficial gray layer and the deeper layers.

Microvascularization of the common tree shrew (Tupaia glis) superior cervical ganglion studied by vascular corrosion cast with scanning electron microscopy.

Microvascularization of the superior cervical ganglion (SCG) of the common tree shrew (Tupaia glis) was investigated by the vascular-corrosion-cast technique in conjunction with scanning electron microscopy. It was found that the SCG of the tree shrew is a highly vascularized organ. It receives arterial blood from branches of the external and common carotid arteries which enter the rostral and caudal portions of the ganglion. These arteries give rise to a subcapsular capillary plexus before branching off to form a group of densely packed intraganglionic capillaries. Moreover, the intraganglionic capillaries tend to follow a tortuous course that is essentially parallel to the longitudinal axis of the ganglion, and they form anastomoses with each other. In addition, the intraganglionic capillaries are also connected to a subcapsular capillary plexus. The capillaries of the SCG converge into venules and collecting veins which subsequently drain rostrally and caudally into the systemic veins. However, neither a pattern of blood vessels resembling glomeruli nor a portal-like intraganglionic microcirculation was observed.

Stereo architecture of the connective tissue cores of the lingual papillae in the treeshrew (Tupaia glis)

The stereo architecture of the lingual connective tissue cores (CTC) in the treeshrew (Tupaia glis) (which has the primitive characteristics of primates) was observed by scanning electron microscopy, and compared to that of other animal orders. The tongue of the treeshrew has three vallate papillae which are situated in the posterior part of the tongue, while some macaques have several vallate papillae. Among numerous filiform papillae, fungiform papillae are sporadically distributed. A filiform papilla consists of a bundle of several slender spine-like processes arranged in a circle at the basal margin. After removal of the epithelium, the CTC of the filiform papilla looks like a human hand raised with the palm facing towards the tongue tip. The fungiform CTC in the threeshrew is columnar in shape (rather similar to that of Insectivora and Rodentia) and at the top there are several round depressions for taste buds. In the treeshrew several large rod-shaped processes are derived from the postero-lateral margin of the tongue, as in Carnivora (dogs and cats), where foliate papillae are located in many other animal species. The treeshrew has numerous characteristics similar to those of the crab-eating macaque (Primates), but at the same time it has some characteristics similar to those of Insectivora, Rodentia, Carnivora and Artiodactyla.

Body temperature rhythm of the tree shrew, Tupaia belangeri.

The circadian rhythm of body temperature of the tree shrew Tupaia belangeri was studied by telemetry. The amplitude of the daily (or circadian) variation was found to be much larger than that of most endotherms (amplitude approximately 5 degrees C) and the bimodal shape of the rhythm differed from the cosine waveform that characterizes the temperature rhythms of most other species. In free-running conditions, as well as in the entrained state, the temperature rhythm remained synchronized to the rhythm of locomotor activity.

Immunohistochemical organization of the ventral lateral geniculate nucleus in the tree shrew.

The ventral lateral geniculate nucleus (vLGN) of the tree shrew (Tupaia belangeri) was differentiated into multiple subdivisions (dorsal cap, intergeniculate leaflet, parvicellular segment, and internal and external magnocellular laminae, the latter being further divisible into a lateral and medial division) on the basis of retinal projections, immunochemistry, and histochemistry. Retinal projections traced with intravitreal injections of wheat germ agglutinin conjugated horseradish peroxidase revealed direct bilateral input to all subregions of the vLGN, except for the internal magnocellular lamina (which received only contralateral input) and the parvicellular segment (which was not retinorecipient). Furthermore, retinal inputs clearly distinguished the relatively heavily retinorecipient intergeniculate leaflet from the less prominently labeled dorsal cap. Immunohistochemical localization of Neuropeptide Y (NPY) perikarya revealed their prominence in the intergeniculate leaflet and the external magnocellular laminae with a concentration along the optic tract. NPY immunoreactive fibers were seen in all but the parvicellular subregion. Gamma amino butyric acid immunoreactivity was seen throughout the vLGN, but was most concentrated in the dorsal cap and the magnocellular laminae, followed by the intergeniculate leaflet. Histochemical studies of cytochrome oxidase and nicotinamide adenosine dinucleotide phosphate (NADPH)-diaphorase localization revealed similar patterns of dense reactivity within the external magnocellular lamina, intergeniculate leaflet and dorsal cap, and somewhat less dense, but substantial reactivity in the internal magnocellular lamina. Within the external magnocellular lamina, cells reactive for cytochrome oxidase were noted in the lateral portion bordering the optic tract, whereas those specific for NADPH-diaphorase were dispersed throughout the lamina. Poor reactivity for both histochemical markers was evident in the parvicellular segment. Overall, the markedly different patterns of retinal input and neurochemical organization between the subdivisions of the tree shrew vLGN suggest their involvement in diverse functions. Furthermore, the basic similarity of the organization of the tree shrew vLGN to that of the taxonomically unrelated ground squirrel may indicate a common mammalian scheme.

Hypophyseal angioarchitecture of common tree shrew (Tupaia glis) revealed by scanning electron microscopy study of vascular corrosion casts

The vascular corrosion cast technique in conjunction with scanning electron microscopy (SEM) was used for the study of pituitary microvascularization in the common tree shrew (Tupaia glis). The pituitary vascular casts were obtained by infusion of low viscosity methyl methacrylate plastic (Batson's no.17) mixture. It was found that the blood supplies to the pituitary complex were from branches of the circle of Willis and could be divided into two groups. The first group consisted of two to four superior hypophyseal arteries (SHAs) branching off from the internal carotid artery supplying each half of the median eminence (ME), infundibular stalk (IS), and pars distalis (PD). The SHAs supplying the ME branched into internal and external capillary plexi. The internal plexus had a larger capillary size (approximately 15 microns in diameter), was deeper in position, and had denser and more complex capillary loops than those in the external plexus. The capillaries of the external plexus were approximately 10 microns in diameter. The two plexi drained into 15-20 hypophyseal portal veins (HPVs) which were located mainly along the ventral and ventrolateral surfaces of the IS before breaking up into large capillaries (approximately 18 microns in diameter) with an anteroposterior arrangement within the PD. The second group consisted of one inferior hypophyseal artery (IHA) on each side branching off from the internal carotid artery. These arteries gave off branches to pierce the dorsolateral and ventrolateral aspects of infundibular process (IP) before branching off to form a capillary network. They also gave rise to radiating capillaries to supply the pars intermedia (PI) surrounding the cortical area of the IP. The hypophyseal cleft separating the PI from the PD was clearly seen with very few blood vessels. The capillaries in both PD and IP joined to form confluent hypophyseal veins draining the blood into the cavernous sinus.

Microvasculature of the Thyroid Gland in the Common Tree Shrew (<i>Tupaia glis</i>): Microvascular Corrosion Cast/Scanning Electron Microscopy Study

A thyroid vascular cast of the common tree shrew (Tupaia glis) was obtained by injection of Batson's No. 17 plastic mixture into the ascending aorta. The cast was studied under the scanning electron microscope. It was found that each half of the gland is supplied by a large superior and a rather small inferior thyroid artery. After plunging into the gland, the arteries divide into smaller branches that are the interlobular, intralobular and follicular arteries (afferent vessels). The basket-like capillaries arising from the follicular arteries and encapsulating thyroid follicles are of large diameter and are arranged in a single layer. The follicular side of the capillary casts was observed to contain numerous small and some large projecting knobs compatible with the presence of fenestrations in the endothelial cells. On the other hand, endothelial nuclear imprints were found mainly on the stromal surface of the follicular capillary casts. Transfollicular capillaries connecting the adjacent follicular capillary networks were also observed. Blood from the follicular capillaries either drains into the follicular veins (efferent vessels) or abruptly drains into the intralobular veins before proceeding to intralobular and interlobular veins, respectively. The interlobular veins are collected into a few small superior, a few larger middle and a few even larger inferior thyroid veins. These veins drain directly into the laryngeal vein lying adjacent to the deep surface of the thyroid gland before joining the jugular vein. Venous valves were identified outside the thyroid gland. In addition, the glomerular capillary island of the parathyroid gland was often seen at the cranioanterolateral and sometimes at the cranioposterolateral aspect of the thyroid gland.

Comparative effects of ouabain on isolated papillary muscle from tree shrews, guinea-pigs and rats

Inotropic effects of ouabain were investigated in the isolated papillary muscle preparations of the tree shrew (Tupaia glis), guinea-pigs and rats. In the guinea-pig and shrew papillary muscles, ouabain at concentrations of 10(-8) to 3 x 10(-7) M caused a concentration-dependent positive inotropic response in a similar magnitude, while in the rat, ouabain at concentrations of 10(-7) to 3 x 10(-6) M elicited negative inotropic one. Either phentolamine or pindolol in a concentration sufficient to block the alpha- and beta-adrenoceptors did not antagonize the positive inotropic effect of ouabain in the papillary muscle preparation of the shrews.

Development of acetylcholinesterase activity in the lateral geniculate nucleus.

The pattern of acetylcholinesterase activity in the tree shrew (Tupaia belangeri) lateral geniculate nucleus (LGN) undergoes a number of striking changes during postnatal development. The adult tree shrew LGN is made up of six cellular layers divided by relatively cell-free interlaminar zones. At birth, however, the nucleus appears unlaminated when processed with conventional Nissl-staining techniques. The cellular lamination appears during the first postnatal week. The eyes open much later, typically at the end of the third week after birth. In the adult tree shrew, acetylcholinesterase (AChE) activity is found throughout the nucleus (both within and between the six cellular layers). In most sections examined, reaction product is slightly more intense in the lateral cell layers (4, 5, and 6). This is in sharp contrast to the pattern at birth (postnatal day zero, or P0). The detectable AChE activity at this age is apparently found in inchoate layers 1-2 and 4-5. Within these pairs, areas innervated by the ipsilateral eye (i.e., incipient layers 1 and 5) appear to contain more reaction product. From P0 to P4, the density of AChE activity increases in layers 1-2 and 4-5 and becomes detectable in the barely evident layers 3 and (usually) 6 at this age. By the middle of the second postnatal week, after laminae are clearly apparent with a Nissl stain, AChE activity has increased and is mainly associated with each cellular layer in the nucleus. During the third week after birth this pattern undergoes a radical shift. The most intense AChE activity is now in the interlaminar zones. Finally, as the adult pattern emerges, AChE activity increases in the cellular layers and all areas of the nucleus exhibit relatively high levels of AChE activity. Superimposed on this changing laminar pattern of AChE activity are changes related to the retinotopic map within the nucleus. Portions of the LGN representing central vision develop their characteristic pattern of activity several days ahead of the regions representing more peripheral visual field locations. AChE activity is also found transiently in the optic tract near the LGN during the first 3 postnatal weeks. Two (possibly three) groups of AChE-carrying fibers can be traced from the optic chiasm to their apparent sites of termination (or origin) in the parabigeminal nucleus, ventral lateral geniculate nucleus, and dorsal LGN. The activity present in the optic tract disappears shortly after eye opening.(ABSTRACT TRUNCATED AT 400 WORDS)

Histochemical localization of cytochrome oxidase activity in the visual system of the tree shrew:normal patterns and the effect of retinal impulse blockage.

The tree shrew Tupaia belangeri has three functional pathways (ON-center, OFF-center, and W-like cells) that arise in the retina and proceed through separate LGN laminae to separate cortical targets. To determine whether these pathways have consistent differences in activity, cytochrome oxidase (C.O.) patterns were examined in the retina, LGN, and striate cortex. In six normal tree shrews the outer and inner plexiform layers of the retina were highly reactive for C.O. A pale, vascularized cleft zone separated the a (OFF) and b (ON) inner plexiform sublaminae, which seemed about equally reactive for C.O. In the LGN, laminae 1 and 2 (ON-center cells) and laminae 4 and 5 (mostly OFF-center cells) were highly reactive for C.O. LGN lamina 3 and 6 are part of an W-like afferent pathway. Lamina 3 was distinctly paler than laminae 1, 2, 4, and 5 while lamina 6 was intermediate. In the striate cortex, layer IV was the most reactive layer. Sublayer IVb (predominantly an OFF region) was consistently more reactive than sublayer IVa (predominantly ON). The middle portion, layer IVm, was paler than either IVa or IVb. This paler region includes, but extends above and below, the cell-sparse "cleft" region. Thus, considering all three levels of the retinogeniculostriate pathway, the ON and OFF systems were equally active until they reached the striate cortex, where the OFF system appeared to be more active than the ON. The W-cell laminae in the LGN exhibited the lowest level of activity. The contribution of ganglion cell activity to these patterns was assessed by intravitreal administration of tetrodotoxin (TTX) blockade either monocularly (three animals) or binocularly (two animals). In the TTX-treated retinae, the inner plexiform a and b sublaminae were paler for C.O., although visible, and were still separated by the pale cleft. The ganglion cell layer was very pale in comparison to the normal. In the LGN, monocular TTX blockade reduced the C.O. reactivity in the ON and OFF laminae that received input from the treated eye but had little effect on the W-like cell laminae. The ipsilaterally innervated ON and OFF laminae were more affected than were the contralaterally innervated laminae. Binocular TTX treatment resulted in a decrease of C.O. activity in the binocular segment of the ON and OFF LGN laminae. In the striate cortex, the most marked changes following TTX treatment occurred in layer IV.(ABSTRACT TRUNCATED AT 400 WORDS)

Laminar organization and ultrastructure of GABA-immunoreactive neurons and processes in the dorsal lateral geniculate nucleus of the tree shrew (Tupaia belangeri).

The distribution and ultrastructure of neurons and neuropil labeled by an antiserum to gamma-aminobutyric acid (GABA) were examined in the lateral geniculate nucleus (LGN) of the tree shrew (Tupaia belangeri). The LGN of this species segregates center type and cell class into three distinct pairs of laminae: a medial pair (laminae 1 and 2) containing ON-center cells, a more lateral pair (4, 5) containing OFF-center cells, and 2 laminae (3, 6) containing W-like cells. The relationship between this laminar segregation and the distribution of GABA immunoreactivity was investigated in the present study. GABA-immunoreactive neurons and neuropil were present in all six of the laminae. However, both the density of labeled cells (adjusted for neuronal density across laminae) and the density of labeled neuropil showed a medial-to-lateral gradient. The adjusted density of labeled cells was higher laterally than medially, and the density of labeled neuropil was significantly greater in the more lateral OFF-center laminae and W-like laminae than in the medial two ON-center laminae. Thus, inhibitory, GABAergic influences may modulate to different degrees the visual signals in the ON, OFF, and W pathways. Labeled cells had a mean cross-sectional area (107 microns 2) approximately one-half that of unlabeled cells (216 microns 2). They constitute 16-34% of the neurons in the LGN. At the electron microscope level, three different kinds of labeled profile were observed. Vesicle containing profiles like the F2 profiles of cat were postsynaptic to retinal terminals and presynaptic to conventional dendrites. F1 axon terminals with dense clusters of vesicles were also labeled as were some myelinated axons. Another labeled profile, which we suggest should be called an F3 process, was a large dendrite of irregular caliber with punctate groups of vesicles near the synapse. Our results suggest that GABAergic circuitry is an important part of the functional organization in the LGN of the tree shrew.

Atrial natriuretic peptide detected by immunocytochemistry in peripheral organs of Tupaia belangeri.

ANP (atrial natriuretic peptide), a peptide found in granules of mammalian atrial cardiac myocytes, has been shown to be active in regulation of blood pressure and body water homeostasis. The existence of ANP in atrium, pituitary, adrenal gland, and kidney of the rat had been immunocytochemically demonstrated with an antibody against rat ANP (102-126). We used the same antibody in immunocytochemical studies for the detection of ANP in peripheral organs of the tree shrew (Tupaia belangeri). The antibody stained granules in myocytes of cardiac atria which indicated that it reacted with tree shrew ANP. In contrast to the rat, no immunoreactive cells were found in pituitaries and adrenal glands. However, in the kidneys distal tubules in outer medulla and cortex were labeled. Ascending limbs of distal tubules were intensely stained when either the peroxidase-antiperoxidase (PAP) or the indirect immunofluorescence method were used. Collecting ducts and convoluted distal tubules in the outer cortex showed a granular type of staining when the immunofluorescence method was used. These data indicate that ANP is present in epithelial cells of distal tubules and collecting ducts, where it may be involved in the regulation of renal salt excretion.

Anatomy and function of the reproductive tract in the captive male tree shrew (Tupaia belangeri).

The reproductive anatomy of the male tree shrew (Tupaia belangeri) was examined and compared with other Tupaiidae. The testes are located prepenially in a pigmented scrotum which is fused to the base of a pendulous penis. The terminal portion of the vas deferens is differentiated into an ampullary gland and joins the duct of the seminal vesicle to form a short ejaculatory duct. The prostate is a compact bilateral body drained by a main collecting duct. In the aggregate, these features indicate that the reproductive system in Tupaia is primate in character. Testicular function in tree shrews is affected by both social and seasonal factors. When males were housed communally, the majority exhibited testicular degeneration accompanied by a loss in the weight and fructose content of the seminal vesicles and in pigmentation of the scrotum. These changes may be due to the presence of dominant conspecifics since animals kept in isolation undergo normal sexual development. Animals captured throughout the year and isolated show seasonal fluctuations in androgenic and spermatogenic function. Reproductive capacity is maximal during the winter and minimal during the summer. Local environmental factors appear to regulate reproductive function so that the greatest number of births occur during the dry season.

An experimental electron microscopical study of a direct retino-pulvinar pathway in the tree shrew

After unilateral enucleation in the tree shrew (Tupaia glis), axo-dendritic synapses degenerate within two secondary visual centers (intergeniculate and lateral pulvinar nuclei). Degenerated optic terminals were found in both pulvinar nuclei on the contralateral side much more often than on the ipsilateral one.

Carcinogenicity of 2,2′-Dihydroxy-di-n-propylnitrosamine in the Tree Shrew (Tupaia glis): Light and Electron Microscopic Features of Pulmonary Adenomas2

The carcinogenic effect of 2,2'-dihydroxy-di-n-propylnitrosamine (DHPN) was examined in the primitive primate, tree shrew (Tupaia glis). DHPN was administered at a dose of 250 mg/kg body weight sc once a week for 80 weeks. Between 65 and 102 weeks, 8 of 9 males given DHPN (89%) and 11 of 14 females given DHPN (78%) developed pulmonary adenomas. In 2 DHPN-treated males, in addition to adenomas, bronchioalveolar carcinomas were observed. Transmission electron microscopic examination of pulmonary adenomas from 4 DHPN-treated animals showed that Clara cells were the main components of these tumors. In addition to pulmonary tumors, 9% of the DHPN-treated animals developed squamous cell carcinomas of the skin and hepatocellular carcinomas. None of the 6 controls, which received olive oil alone, developed any tumors.

Transneuronal transport of tritiated fucose and proline in the visual pathways of tree shrew Tupaia glis

Transneuronal transport of tritiated fucose and proline in the visual pathways of tree shrew Tupaia glis

Organization of the visual afferents into the LGd and the pulvinar of the tree shrew Tupaia glis

Organization of the visual afferents into the LGd and the pulvinar of the tree shrew Tupaia glis

Temperature regulation in the tree shrew Tupaia glis

1. 1. The mean day-night difference of body temperature in Tupaia glisis5-1°C. 2. 2. The nocturnal standard metabolic rate is 0.76 ml O 2/g per hr ± 0.07 (2 S.E.) at a mean body temperature of 37.0°C. 3. 3. The thermal neutral zone extends from near 30 to above 37°C. 4. 4. The marked decline in body temperature at lower ambient temperatures is interpreted to reflect the tropical habitation of this species.