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The tree shrews: adjuncts and alternatives to primates as models for biomedical research.

The tree shrews are non-rodent, primate-like, small animals. There is increasing interest in using them to establish animal models for medical and biological research. This review focuses on the use of the tree shrews in in vivo studies on viral hepatitis, hepatocellular carcinoma (HCC), myopia, and psychosocial stress. Because of the susceptibility of the tree shrews (Tupaia belangeri) and their hepatocytes to infection with human hepatitis B virus (HBV) in vivo and in vitro, these animals have been used to establish human hepatitis virus-induced hepatitis and human HBV- and aflatoxin B1-associated HCC models. As these animals are phylogenetically close to primates in evolution and have a well-developed visual system and color vision in some species, they have been utilized to establish myopia models. Because dramatic behavioral, physiological, and neuroendocrine changes in subordinate male tree shrews are similar to those observed in depressed human patients, the tree shrews have been successfully employed to experimentally study psychosocial stress. However, the tree shrews holds significant promise as research models and great use could be made of these animals in biomedical research.

Animals↗

Postnatal growth pattern of F1 hybrids of a cross between two strains of large and small musk shrews, Suncus murinus.

In our previous comparison of the postnatal growth of two laboratory strains of the musk shrew (Suncus murinus), the large shrews originating in Bangladesh (BAN strain) and the small shrews from Nagasaki, Japan (NAG strain), we found that differences in the 120-day adult body weight of the two strains (BAN: males 135.3g and females 82.0g; NAG: 52.9 and 34.2g, respectively) reflect both an approximately 2.5 times longer linear growth phase and an approximately 1.5 times higher growth rate during this phase in the BAN strain. We examined the postnatal growth (from 5 to 120 days after birth) of the F1 hybrids of a cross between the two strains and compared our findings with the above-mentioned growth data for the parent strains. A growth equation composed of linear and monomolecular functions was fitted to the growth data for the shrews. The growth trajectories of both sexes of the F1 hybrids were situated roughly midway between those of the two parent strains throughout the entire growth process. The adult body weight of the F1 shrews at 120 days of age averaged 86.0g in the males and 51.7g in the females. The growth phase lasted approximately 60 days in the males and approximately 40 days in the females of both the F1 shrews and the parent strains. The duration of the linear growth phase of the F1 hybrids (29.8 days in the males and 19.0 days in the females), however, was approximately intermediate between the parent strains (BAN: males 33.4 days and females 26.1 days; NAG: 18.8 and 13.9 days, respectively).(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Induced myopia associated with increased scleral creep in chick and tree shrew eyes.

PURPOSE: To investigate the role of scleral creep in the axial elongation of chick and tree shrew eyes with induced myopia. METHODS: Form-deprivation myopia was induced with a diffusing occluder worn over one eye. Scleral samples from the posterior pole and equatorial regions of myopic, contralateral (control), and age-matched normal chick and tree shrew eyes were loaded in vitro with a force of 5 g for 20 minutes while creep extension was monitored. The elastic behavior of sclera from myopic, control, and normal chick eyes was also compared. RESULTS: In both chick and tree shrew, posterior and equatorial scleral samples from myopic eyes had significantly (P < 0.05) greater creep extensions than equivalent samples from control and normal eyes (n = 10, each group). Among individual tree shrews the difference in creep rate between the sample from the myopic eye and that from the control eye correlated with vitreous chamber elongation (r = 0.746, P < 0.05) and development of myopia (r = 0.792, P < 0.01) in the deprived eye. No such association was found in the data from chicks. The elastic properties of chick sclera were unaffected in form-deprivation myopia. CONCLUSIONS: In chick and tree shrew, form-deprivation myopia is associated with increased creep rate of posterior and equatorial sclera. In tree shrew, the correlation between increased scleral creep rate and vitreous chamber elongation in myopic eyes supports the hypothesis that induced changes in the axial length of the mammalian eye are mediated by changes in the creep properties of the sclera.

Animals↗

[Alteration of p53 gene during tree shrews' hepatocarcinogenesis].

OBJECTIVE: To detect the expression and variation of p53 gene during tree shrews' hepatocarcinogenesis induced by hepatitis B virus (HBV) and aflatoxin B1 (AFB1). METHODS: Tree shrews were divided into four groups: the tree shrews were infected with HBV and fed with AFB1 in group A, only infected with HBV in group B, fed with AFB1 alone in group C, and normal control in group D. All the tree shrews were performed liver biopsy every 15 weeks. The tissues of liver and tumor were detected by immunohistochemistry and molecular biotechnologies. RESULTS: (1) The incidence of hepatocellular carcinoma (HCC) in group A (66.7%) was higher than that in Group B and C (30%). HCC appearance in group A was earlier than that in group C (120.0 weeks +/-16.6 weeks vs 153.3 weeks +/-5.8 weeks, t = 3.336, P<0.01). (2) Mutated p53 protein was not found before the 75th week of the experiment in each group. (3) At the 105th week, the expression rates of mutated p53 protein were 78.6%, 60% and 71.4% in group A, B and C respectively, which were much higher than that (10%) in group D (x2 > or = 5.03, P<0.05). An abnormal band of p53 gene was detected in both group A and C. (4) The mutation points of p53 gene in liver cancer of tree shrew were at codon 275, 78 and 13. The nucleotide sequence and amino acids sequence of tree shrew's wild-type p53 showed 91.7% and 93.4% homology with those of human p53 respectively. CONCLUSIONS: There is a remarkable synergistic effect between HBV and AFB1 on HCC. Mutated p53 protein is expressed before HCC occurrence, which promotes the development and progress of HCC. HBV and AFB1 may synergistically induce p53 gene mutation.

Aflatoxin B1↗

A shrew-sized origin for primates.

The origin of primates has had a long history of discussion and debate, with few ever considering the impact of the original body weight on subsequent primate adaptive radiations. Here, I attempt to reconstruct early primate evolution by considering the initial size of primates as well as the critical functional-adaptive events that had to occur prior to the early Eocene. Microcebus is often viewed as a living model, and thus 40-65 g might represent a practical ancestral weight for the origin of primates. I consider a smaller original body weight, likely 10-15 g in actual size, and I address the biological implications for shrew-sized primates by comparing the behavioral ecology of mouse lemurs, our smallest living primates, to another tiny-sized mammalian group, the shrews (Family Soricidae). Several behavioral and ecological characteristics are shared by shrews and mouse lemurs, and several mammalian trends are evident with decreased size. I suggest that a shrew-sized ancestral primate would have had high metabolic, reproductive, and predation rates, relatively low population densities, and a dispersed and solitary existence with a promiscuous mating system. Although small mammals like shrews provide insights concerning the ancestral size of primates, primate origins have always been tied to arboreality. I assess other potential arboreal models such as Ptilocercus and Caluromys. By combining all of this information, I try to sequence the events in a functional-adaptive series that had to occur before the early Eocene primate radiations. I suggest that all of these important adaptive events had to occur at a small body size below 50 g.

Adaptation, Physiological↗

Organization of the somatosensory cortex in elephant shrews (E. edwardii).

The superorder Afrotheria consists of a diverse group of mammals, including elephants, hyraxes, dugongs, sea cows, aardvarks, tenrecs, golden moles, and elephant shrews. Recent studies suggest this clade diverged from other placental mammals 100 million years ago and thus may represent the sister group to the remaining placental mammals. Despite this important taxonomic position, relatively few studies have investigated cortical organization in these species. Here we present results of an investigation of the somatosensory cortex in the Cape elephant shrew (Elephantulus edwardii). Using multiunit electrophysiological recording techniques, we identified a topographic map of the elephant shrew's body in a location and orientation consistent with the primary somatosensory cortex (S1). The elephant shrew's elongated snout, extensive facial vibrissae, and long tongue accounted for a large portion of the somatosensory representation, located in a relatively rostral area of cortex. Evidence for an additional somatosensory area, presumed to be secondary somatosensory cortex (S2), was found just lateral to S1. Visual and auditory responsive areas were also identified and the extent of visual cortex appeared to be quite large in these highly visual mammals. Despite the elephant shrew's exceptionally well-developed eyes, ears, and vibrissae, there were no anatomical correlates to sensory areas, or body part representations (e.g., barrels), that could be identified in the flatted cortex.

Animals↗

The musk shrew, Suncus murinus, a unique animal model for the study of female behavioral endocrinology.

Insectivora is the third largest mammalian order, composed of a number of unusual species considered to be the most primitive of the modern eutherian mammals. Yet, little is known about the members of this evolutionarily unique and important group. The musk shrew (Suncus murinus) is a convenient and practical laboratory animal. The study of the reproductive biology of this species will yield needed comparative data. Moreover, the little information that we have collected on this species suggests that several unusual characteristics make this animal a worthwhile and novel model for endocrine research. Most of the current and past research on this species has focused on the endocrinology of the female musk shrew. Unlike conventional small mammal models, the female musk shrew has no spontaneous ovarian cycle. The ovaries of the adult, unmated female do not undergo spontaneous follicular development. At the time of mating only small, relatively immature follicles are present. As a consequence, ovarian hormone production is not cyclic. Thus, traditional hormone target tissues such as the vaginal epithelium and the uterus do not exhibit cycles in cell proliferation. Sexual behavior is likewise demonstrated in an acyclic manner. Virgin females become sexually receptive within minutes after their first exposure to a male, and nonpregnant females are virtually always sexually receptive. Sexual receptivity in the virgin female musk shrew occurs in the face of relatively low plasma estradiol levels and higher androgen levels, and does not appear to be mediated via ovarian estradiol. Instead, recent work suggests that close to physiological doses of testosterone can restore sexual behavior in ovariectomized musk shrews.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Modelling and monitoring organochlorine and heavy metal accumulation in soils, earthworms, and shrews in Rhine-delta floodplains.

In the Rhine-delta, accumulation of microcontaminants in floodplain foodwebs has received little attention in comparison with aquatic communities. To investigate organochlorine and metal concentrations in a terrestrial foodchain, samples of soil, earthworms (Lumbricus rubellus), and shrew (Crocidura russula, Sorex araneus) livers and kidneys were taken from two moderately to heavily polluted floodplains. Chlorobiphenyl residues in earthworm fat were 0.10 to 3.5 times the concentrations in soil organic matter, whereas ratios for other organochlorines varied between 0.87 and 8.8. These ratios are one order of magnitude lower than expected from laboratory experiments with earthworms, and laboratory and field studies on aquatic invertebrates. Bioconcentration ratios for heavy metals are in accordance with literature values for other locations, confirming the high potential for cadmium accumulation in Lumbricidae. Concentrations of organochlorines in shrew liver lipids were 1.0 to 13 times the residues in earthworm fat. These values are higher than lipid-corrected biomagnification ratios for laboratory rodents, but equal to those measured for benthivorous birds in the Rhine-delta. On a dry weight basis, kidney-earthworm ratios for cadmium were about one order of magnitude lower than previously reported values for insectivores. Soil concentrations of many compounds in both floodplains did not meet Dutch quality standards. Yet, hexachlorobenzene, chlorobiphenyl 153 (PCB153), gamma-hexachlorocyclohexane, sigma DDT, and dieldrin residues in earthworms and shrews did not exceed diet levels expected to be safe for endothermic species. An exception was noted for cadmium in worms and shrew kidneys. Heavy metal pollution in soil was close to levels that are critical to earthworms in laboratory studies. Cadmium concentrations in shrew kidneys were below levels suggested to be safe for Sorex araneus, but above those that were critical to the rat.

Animals↗

A comparative analysis of the potential of cannabinoids and ondansetron to suppress cisplatin-induced emesis in the Suncus murinus (house musk shrew).

RATIONALE: The 5-HT3 antagonist, ondansetron (OND), and the cannabinoid, delta9-tetrahydrocannabinol (delta9-THC), have been shown to interfere with emesis; however, their relative and/or combined effectiveness in suppressing vomiting produced by the chemotherapeutic agent, cisplatin, is unknown. OBJECTIVE: To evaluate the potential of: 1) a broad range of doses of delta9-THC and OND to prevent cisplatin-induced vomiting and retching in the Suncus murinus (house musk shrew), 2) combined treatment with ineffective individual doses of delta9-THC and OND to prevent cisplatin-induced vomiting and retching, 3) the CB1 receptor antagonist, SR141716, to reverse the antiemetic effects of OND, and 4) cannabidiol (CBD), the principal non-psychoactive component of marijuana, to reverse cisplatin-induced vomiting in the shrew. METHODS: Shrews were injected with various doses of OND (0.02-6.0 mg/kg), delta9-THC (1.25-10 mg/kg) and a combination of ineffective doses of each (0.02 mg/kg OND+1.25 mg/kg delta9-THC) prior to being injected with cisplatin (20 mg/kg) which induces vomiting. Shrews were also injected with CBD (5 mg/kg and 40 mg/kg) prior to an injection of cisplatin. RESULTS: OND and delta9-THC both dose-dependently suppressed cisplatin-induced vomiting and retching. Furthermore, a combined pretreatment of doses of the two drugs that were ineffective alone completely suppressed vomiting and retching. CBD produced a biphasic effect, suppressing vomiting at 5 mg/kg and potentiating it at 40 mg/kg. CONCLUSIONS: A low dose of the non-intoxicating cannabinoid CBD may be an effective anti-emetic treatment and combined doses of OND and delta9-THC that are ineffective alone suppresses cisplatin-induced emetic reactions in shrews.

Animals↗

Magnificent expression of asialo GM1 on thymus cells and spleen T cells of the musk shrew, Suncus murinus.

The expression of asialo GM1 (GA1) was observed on almost all thymocytes from young musk shrew, at the age of 4 weeks by flow cytometric analysis. In adult shrew aged 10 months, the ratio of GA1-negative thymocytes was increased. Among several anti-glycolipid antibodies used, anti-GM1 and anti-Forssman also reacted with the thymocytes. Protein fraction of the thymocytes was analyzed by SDS-PAGE followed by immunoblotting. Anti-GA1 and anti-GM1 showed two bands and one band, respectively, however, their mobilities were different from each other. Anti-Forssman did not stain any protein. The GA1-positive population in spleen T cell fraction was not detected in young shrew but most of the T cells were changed to GA1-positive cells in adult shrew. When mixed lymphocyte culture was performed, the GA1-negative spleen T cells in young shrew were changed to express GA1 marker on their cell surface by differentiation. Abbreviations used were as follows: GA1, Gal beta 1-3GalNAc beta 1-4Gal beta 1-4Glc-Cer; GM1, Gal beta 1-3GalNAc beta 1-4(NeuAc alpha 2-3)Gal beta 1-4Glc-Cer; Forssman, GalNAc alpha 1-3GalNAc beta 1-3Gal alpha 1-4Gal beta 1-4Glc-Cer.

Animals↗

Intestinal disaccharidases in the house musk shrew, Suncus murinus: occurrence of sucrase deficiency.

1. Disaccharidase activities of the small-intestinal brush border membrane were studied in six laboratory lines of the house musk shrew, Suncus murinus. 2. Sucrase activity was detected in all shrews of one line, but not in any shrew of three lines. In the other two lines it was found in some shrews, but not in the others. 3. Maltase, isomaltase, trehalase and lactase activities were found in all shrews of all the lines examined. 4. Sucrase was normally associated with isomaltase to form an enzyme complex. 5. Detergent-solubilized isomaltase, whether associated with sucrase or not, was inhibited by antibodies against rabbit sucrase-isomaltase to almost the same extent as the rabbit one, suggesting that isomaltase is not affected by a mutation(s) in sucrase.

Animals↗

Metal bioaccumulation in the greater white-toothed shrew, Crocidura russula, inhabiting an abandoned pyrite mine site.

Hepatic and renal concentrations of iron, magnesium, zinc, lead, copper, manganese, mercury, cadmium, molybdenum, chromium, and nickel were quantified in shrews (Crocidura russula) inhabiting a pyrite mine site in Portugal. Several morphological parameters (body weight, residual index, and relative weights) were also examined to clarify the physiological effects of pollution. Shrews from the mine showed increased bioavailability of Fe, Pb, Hg, Cd, Mo, and Ni in comparison with reference specimens. Adult shrews had the highest Cd levels while Cr and Ni concentrations diminished. Intersexual differences were found for Mo and Ni. As a consequence of metal pollution, the relative hepatic weight was higher in shrews from the mine site when compared with reference specimens. These data indicate that C. russula is a good bioindicator of metal pollution. We also evaluated the toxic effects of Pb, Hg, Cd, and Ni, because several shrews from the polluted site showed high concentrations of these metals. To approximate at the real biological impact of abandoned mines, after this first step it is necessary to associate the bioaccumulation levels and morphological effects with other physiological, ecological and genetical biomarkers.

Age Factors↗

Localization of nitric oxide synthase in the tree shrew retina.

Nitric oxide (NO) is a novel neuronal messenger that likely influences retinal function by activating retinal guanylyl cyclase to increase levels of cGMP. In the present study, the localization of neuronal nitric oxide synthase (nNOS, Type I NOS) in the cone-dominant tree shrew retina was studied using NADPH-d histochemistry and nNOS immunocytochemistry. Both NADPH-d and nNOS-immunoreactivity (IR) labeled the inner segments of rods and the myoids of a regular subpopulation of cones, with their corresponding nuclei outlined. The labeled cone myoids were co-localized with a marker for short-wave-sensitive (SWS) cones (S-antigen) and also displayed the regular triangular packing and density (7%) characteristic of SWS cones in tree shrew and other mammalian retinas. These measures confirmed the identity of the labeled cones as SWS cones. Photoreceptor ellipsoids of all cones were strongly labeled by NADPH-d reactivity, but lacked nNOS-IR. Another novel finding in tree shrew retina was that both NADPH-d and nNOS-IR labeled Muller cells, which have not been labeled by nNOS-IR in other mammalian retinas. Consistent with findings in rod-dominant retinas, two types of amacrine cells at the vitreal edge of the inner nuclear layer and a subpopulation of displaced amacrine cells at the scleral edge of the ganglion cell layer were labeled by both NADPH-d and nNOS-IR. Processes of these labeled cells were seen to extend into the inner plexiform layer, where dense punctate label was seen, especially in the central sublamina. These results show that localization of NOS in the cone-dominant tree shrew retina shares some common properties with rod-dominant mammalian retinas, but also shows some species-specific characteristics. The new finding of nNOS localization in tree shrew SWS cones and rods, but not in other cones, raises interesting questions about the roles of NO in the earliest level of visual processing.

Animals↗

Novelties of conception in insectivorous mammals (Lipotyphla), particularly shrews.

In the order Lipotyphla (Insectivora), certain reproductive features differ quite distinctly from the eutherian norms, and are of interest with regard to the evolution of mammalian gamete function and perhaps for questions of lipotyphlan phylogeny. As seen in one or more members of five lipotyphlan families (shrews, moles, hedgehogs, golden moles, tenrecs), these features can involve the configuration of the male tract including the penis, the morphology of the sperm head, the anatomy of the oviduct and the patterns of sperm transport within it, the character of the cumulus oophorus, and the way in which fertilising spermatozoa interact with the eggs. However, the picture is by no means uniform within the order. Reproductive idiosyncrasies occur variously in the different lipotyphlan families, and appear consistently and strikingly in shrews--the group studied most extensively. Compared to the patterns in most Eutheria, the most interesting anomalies in soricids include (a) the regulation of sperm transport to the site of fertilisation by oviduct crypts, whose arrangement can vary even according to species, (b) a circumscribed matrix-free cumulus oophorus that appears essential for fertilisation as the inducer of the acrosome reaction, (c) barbs on the acrosome-reacted sperm head by which it may attach to the zona pellucida. With regard to the bearing such reproductive traits might have on lipotyphlan systematics, the African mouse shrew (Myosorex varius) displays a mix of traits that characterize either crocidurine or soricine shrews, consistent with the proposal that it belongs in a more primitive tribe, Myosoricinae, or subfamily, the Crocidosoricinae, from which the crocidurine and soricine lines probably evolved. Moreover, although elephant shrews are assigned now to a separate order (Macroscelidea), they display several of the unusual reproductive features seen in lipotyphlans, particularly in chrysochlorids and tenrecs. On the other hand, if used as a phylogenetic yardstick, none of the reproductive features described serves to define the Lipotyphla as classically constituted within one order, nor necessarily all the relationships suggested by recent sequencing studies of nuclear and mitochondrial genes.

Animals↗

Attempted isolation of Blastomyces dermatitidis from native shrews in northern Wisconsin, USA.

The precise ecological niche of Blastomyces dermatitidis is unknown. The related dimorphic fungus, Paracoccidioides brasiliensis, has been isolated from South American ground-dwelling insectivorous armadillos. We attempted to isolate Blastomyces from shrews, North American ground-dwelling insectivores that have been shown to harbor Histoplasma capsulatum in endemic areas. Forty-seven masked shrews (Sorex cinereus) and 13 northern short-tailed shrews (Blarina brevicauda) were collected in endemic areas of northern Wisconsin and Michigan using pitfall traps. Specimens were collected between 1998 and summer 2002, stored frozen, then necropsied. Cultures of nasopharynx, lungs, liver, spleen and large and small bowel were placed on yeast extract phosphate agar with one or two drops of ammonium hydroxide. Cultures for Blastomyces were negative from all 60 shrews and two deer mice (Peromyscus maniculatus) and three southern red-backed voles (Clethrionomys gapperi), which were trapped inadvertently. Histological examination of 36 of these specimens revealed no Blastomyces yeast forms. Northern Wisconsin shrews do not appear to be carriers of B. dermatitidis.

Animals↗

Age-associated and cell-type specific changes in NGF requirement for neurite regeneration from trigeminal ganglion cells of the shrew (Suncus murinus).

Low density (20-50 cells/cm2), dissociated cultures of trigeminal ganglion (TRG) cells of the shrew, were made in a serum-free medium, and neurite growth was compared across ages of the animal and between NGF-free and NGF-rich conditions. TRG cells from newborn shrew (0-3 days old), which grew long neurites in an NGF-rich medium, failed to grow any neurites in an NGF-free medium. In contrast, TRG cells from aged shrew (16-19 months), which grew neurites without NGF, exhibited no further increment in neurite length when NGF (50-200 ng/ml) was added to the culture medium. TRG cells from adult shrew (4-5 months) were revealed to be a mixture of NGF-dependent and NGF-independent cells. The NGF-dependent cells (20% of the population) had large-sized somata of 24-32 microns diameter (L-type cells) and 2-7 long neurites enriched with arborizations. The remaining NGF-independent cells (80%) had small-sized somata (15-25 microns, S-type cells), and grew 1-3 neurites with a small number of arborizations. These findings suggest that the primary sensory neuron of the shrew has a cell-type specific critical period in the aging process with respect to the requirement of NGF for neurite promotion.

Aging↗

Chromosome location of sixteen genes in the common shrew, Sorex araneus L. (Mammalia, Insectivora).

This report extends the genetic map of the common shrew (Sorex araneus) by use of a clone panel of shrew-Chinese hamster and shrew-mouse hybrid cells (Pack et al., 1995; Matyakhina et al., 1996). This set of hybrid clones made it possible to assign the shrew genes for isocitrate dehydrogenase 2 (IDH2), inorganic pyrophosphatase (PP), glutamicpyruvate transaminase (GPT), adenosine kinase (ADK), glucuronidase 2 (GUSB) and acid phosphatase 2 (ACP2) to chromosome ik; the genes for adenylate kinases 1 and 3 (AK1 and AK3) to chromosome af; the genes for glutamate-oxaloacetate transaminase 2 (GOT2), peptidase D (PEPD) and growth hormone (GH) to chromosome hn; the gene for phosphoglucomutase 2 (PGM2) to chromosome go, the gene for enolase 1 (ENO1) to chromosome ji, the gene for ornithine carbamoyl-transferase (OTC) to chromosome de, the gene for aminoacylase 1 (ACY1) to arm m (chromosome mp), the gene for glutamate-oxaloacetate transaminase 1 (GOT1) to arm q (chromosome qr). Thus, the genetic map of the common shrew now contains 33 genes and it is possible to compare the syntenic associations with other species.

Animals↗

Localization of aquaporin water channels in the airway of the musk shrew (Suncus murinus) and the rat.

Aquaporins (AQPs) constitute a family of water channels that facilitate membrane water permeability in various tissues of animals. In this study, we compared the expression and localization of AQPs in the respiratory system of the musk shrew (Suncus murinus), which is an insectivore, and the rat by immunohistochemical methods. In both the musk shrew and the rat, AQP1 was expressed throughout the airway in endothelial cells of subepithelial blood vessels and in nasal submucosal fibroblasts. AQP3 and AQP4 were detected in neither the epithelium nor the subepithelial layer of the musk shrew airway, but were abundant in the rat airway epithelium. Musk shrew AQP5 was distributed in the superficial epithelial cells facing the airspaces and in submucosal glandular cells, but, unlike in the rat, not in lung alveolar cells. Additionally, the expression patterns of AQP4 and AQP5 of the musk shrew were partly similar to those of the human previously reported, absence of AQP4 and presence of AQP5 in the upper airway. The expression differences of AQPs between species in the airway indicate that the physiological importance of each AQP may be different in each species.

Animals↗