著者

Jun IWAMOTO Tsuyoshi TAKEDA Yoshihiro SATO

出版者

Japanese Association for Laboratory Animal Science

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

vol.54, no.1, pp.1-6, 2005 (Released:2005-02-23)

参考文献数

31

被引用文献数

55 71

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Increasing peak bone mass at skeletal maturity, minimizing bone loss during middle age and after menopause, and increasing bone mass and preventing falls in advanced age are important measures for preventing osteoporotic fractures in women. Exercise has generally been considered to have a positive influence on bone health. This paper reviews the effects of treadmill exercise on bone in young, adult, ovariectomized, and osteopenic female rats. Treadmill exercise increases cortical and cancellous bone mass of the tibia as a result of increased bone formation and decreased bone resorption in young and adult rats. The increase in lumbar bone mass seems to be more significant when long-term exercise is applied. Treadmill exercise prevents cancellous bone loss at the tibia as a result of suppressed bone resorption in ovariectomized rats, and increases bone mass of the tibia and mechanical strength of the femur, as a result of suppressed bone resorption and increased bone formation in osteopenic rats after ovariectomy. Treadmill exercise transiently decreases the serum calcium level as a result of accumulation of calcium in bone, resulting in an increase in serum 1,25-dihydroxyvitamin D3 level and a decrease in serum parathyroid hormone level. We conclude that treadmill exercise may be useful to increase bone mass in young and adult rats, prevent bone loss in ovariectomized rats, and increase bone mass and bone strength in osteopenic rats, especially in the long bones at weight-bearing sites. Treadmill exercise may have a positive effect on the skeleton in young, and adult, ovariectomized, and osteopenic female rats.

著者

Yui SHIMOMI Yasuhiko KONDO

出版者

Japanese Association for Laboratory Animal Science

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

pp.19-0161, (Released:2020-06-30)

被引用文献数

3

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Among the intact male rats, a subpopulation has been found to show little or no sexual behavior, even after experiencing several mating sessions. This study investigated whether sexually sluggish (SS) males show behavioral differences from normal copulatory (NC) males, other than those concerning sexual behavior. The olfactory preference of males was measured through the time spent displaying nose-poking behavior directed at sexually active males and estrous females for odor exploration in a three-chamber apparatus. Both the NC and SS males showed a significant preference for the odor of estrous females compared with that of male odors. However, SS males spent significantly less time nose-poking estrous females than NC males. The food-finding test was performed after overnight fasting. Our findings showed that all the NC males found the buried pellet within 5 min, whereas over 60% of the SS males failed to find it. The males were also tested for their ability to find a buried bag containing soiled bedding from estrous female cages. The bag was found by 80% of NC males, but only by 20% of SS males. Our results suggest that SS and NC male rats differ not only in sexual behavior but also in other functions such as olfaction.

著者

坪井 基枝 林 良敬 城ヶ原 貴通 小倉 剛 村田 善晴 織田 銑一

出版者

Japanese Association for Laboratory Animal Science

雑誌

Experimental animals (ISSN:13411357)

巻号頁・発行日

vol.58, no.1, pp.31-40, 2009-01-01

著者

Suvd BYAMBAA Hideki UOSAKI Hiromasa HARA Yasumitsu NAGAO Tomoyuki ABE Hiroaki SHIBATA Osamu NUREKI Tsukasa OHMORI Yutaka HANAZONO

出版者

Japanese Association for Laboratory Animal Science

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

pp.19-0120, (Released:2019-12-04)

被引用文献数

4

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X-linked severe combined immunodeficiency (X-SCID) is an inherited genetic disorder. A majority of X-SCID subjects carries point mutations in the Interleukin-2 receptor gamma chain (IL2RG) gene. In contrast, Il2rg-knockout mice recapitulating X-SCID phenotype lack a large part of Il2rg instead of point mutations. In this study, we generated novel X-SCID mouse strains with small insertion and deletion (InDel) mutations in Il2rg by using Clustered Regularly Interspaced Short Palindromic Repeats (CRISPR)/Cas9. To this end, we injected Streptococcus pyogenes Cas9 (SpCas9) mRNA and single guide RNA targeting the exon 2, 3 or 4 of Il2rg into mouse zygotes. In the F0 generation, we obtained 35 pups and 25 out of them were positive for Surveyor assay, and most of mutants displayed dramatic reductions of T and B lymphocytes in the peripheral blood. By amplicon sequencing, 15 out of 31 founder mice were determined as monoallelic mutants with possible minor mosaicisms while 10 mice were mosaic. Finally, we established new strains with 7-nucleotide deletion and 1-nucleotide insertions in the exon 2 and the exons 3 and 4, respectively. Although no IL2RG protein was detected on T cells of exons 3 and 4 mutants, IL2RG protein was unexpectedly detected in the exon 2 mutants. These data indicated that CRISPR/Cas9 targeting Il2rg causes InDel mutations effectively and generates genetically X-SCID mice. Genetic mutations, however, did not necessarily grant phenotypical alteration, which requires an intensive analysis after establishing a strain to confirm their phenotypes.

著者

Yu-Hsin CHANG Megumi C. KATOH Ahmed M. ABDELLATIF Guli XIAFUKAITI Abdelaziz ELZEFTAWY Masami OJIMA Seiya MIZUNO Akihiro KUNO Satoru TAKAHASHI

出版者

Japanese Association for Laboratory Animal Science

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

pp.19-0105, (Released:2019-12-02)

被引用文献数

1

---

Cre/loxP is a site-specific recombination system extensively used to enable the conditional deletion or activation of target genes in a spatial- and/or temporal-specific manner. A number of pancreatic-specific Cre driver mouse lines have been broadly established for studying the development, function and pathology of pancreatic cells. However, only a few models are currently available for glucagon-producing α-cells. Disagreement exists over the role of the MAFB transcription factor in glucagon expression during postnatal life, which might be due to the lack of α-cell-specific Cre driver mice. In the present study, we established a novel Gcg-Cre knock-in mouse line with the Cre transgene expressed under the control of the preproglucagon (Gcg) promoter without disrupting the endogenous Gcg gene expression. Then, we applied this newly developed Gcg-Cre mouse line to generate a new α-cell-specific Mafb conditional knockout mouse model (MafbΔGcg). Not only α-cell number but also glucagon production were significantly decreased in MafbΔGcg mice compared to control littermates, suggesting an indispensable role of MAFB in both α-cell development and function. Taken together, our newly developed Gcg-Cre mouse line, which was successfully utilized to uncover the role of MAFB in α-cells, is a useful tool for genetic manipulation in pancreatic α-cells, providing a new platform for future studies in this field.

著者

Qing-Shan GAO Mei-Fu XUAN Zhao-Bo LUO Hyo-Jin PAEK Jin-Dan KANG Xi-Jun YIN

出版者

Japanese Association for Laboratory Animal Science

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

pp.19-0018, (Released:2019-07-15)

被引用文献数

3

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The nuclear receptor corepressor Hairless (HR) interacts with nuclear receptors and controls expression of specific target genes involved in hair morphogenesis and hair follicle cycling. Patients with HR gene mutations exhibit atrichia, and in rare cases, immunodeficiency. Pigs with HR gene mutations may provide a useful model for developing therapeutic strategies because pigs are highly similar to humans in terms of anatomy, genetics, and physiology. The present study aimed to knockout the HR gene in pigs using the clustered regularly interspaced short palindromic repeat (CRISPR)/CRISPR-associated-9 system and to investigate the molecular and structural alterations in the skin and thymus. We introduced a biallelic mutation into the HR gene in porcine fetal fibroblasts and generated nine piglets via somatic cell nuclear transfer. These piglets exhibited a lack of hair on the eyelids, abnormalities in the thymus and peripheral blood, and altered expression of several signaling factors regulated by HR. Our results indicate that introduction of the biallelic mutation successfully knocked out the HR gene, resulting in several molecular and structural changes in the skin and thymus. These pigs will provide a useful model for studying human hair disorders associated with HR gene mutations and the underlying molecular mechanisms.

著者

Saki TOME Tetsuya SASAKI Satoru TAKAHASHI Yosuke TAKEI

出版者

Japanese Association for Laboratory Animal Science

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

pp.19-0038, (Released:2019-06-10)

被引用文献数

3

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T helper 17 (Th17) cells have been suggested to play a crucial role in various complications during pregnancy by participating in maternal immune activation (MIA). To test a possible role for Th17 cells in MIA-mediated abortion, we analyzed transgenic mice overexpressing retinoic acid-related orphan receptor gamma-t (RORγt), a master regulator of IL-17 producing cell development. These mutant mice (RORγt Tg mice) exhibited a constitutive upregulation of serum interleukin-17A (IL-17A) and decreased E-cadherin expression in cell–cell junctions of placental tissues. Abortion after the administration of a viral-mimicking synthetic double-stranded RNA polyinosinic–polycytidylic acid was more frequent in RORγt Tg mice than wild-type mice. These results suggest that excessive Th17 cell activity alters immune responsiveness and increases the rate of abortion during gestation.

著者

Shayida MAIMAITI Ryusuke KOSHIDA Masami OJIMA Kaushalya KULATHUNGA Hisashi OISHI Satoru TAKAHASHI

出版者

Japanese Association for Laboratory Animal Science

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

pp.18-0182, (Released:2019-05-16)

被引用文献数

3

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Mammalian postnatal growth is regulated primarily by the growth hormone (GH)/insulin-like growth factor I (IGF-I) axis. MafB is a basic leucine zipper (bZip) transcription factor that has pleiotropic functions. Although MafB plays a critical role in fetal brain development, such as in guidance for hindbrain segmentation, its postnatal role in neurons remains to be elucidated. To investigate this, we used neuron-specific Mafb conditional knockout (cKO) mice. In addition to an approximately 50% neonatal viability, the Mafb cKO mice exhibited growth retardation without apparent signs of low energy intake. Notably, serum IGF-I levels of these mice in the postnatal stage were lower than those of control mice. They seemed to have a neuroendocrine dysregulation, as shown by the upregulation of serum GH levels in the resting state and an inconsistent secretory response of GH upon administration of growth hormone-releasing hormone. These findings reveal that neuronal MafB plays an important role in postnatal development regulated by the GH/IGF-I axis.

著者

Shunichi SUZUKI Daiichiro FUCHIMOTO

出版者

Japanese Association for Laboratory Animal Science

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

pp.18-0142, (Released:2019-01-14)

被引用文献数

4

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Tonsils are mucosa-associated lymphoid tissues located at the openings of the gastrointestinal and respiratory tracts, which play a key role in the surveillance of inhaled or ingested pathogens and can concurrently be reservoirs of infectious agents. Therefore, tonsils are important for the immunology and hygiene management of domestic animals, including pigs. However, the process of their fetal developmental has been poorly described, at least in part, because rodents lack tonsils. Therefore, we performed a histological analysis of porcine tonsils of the soft palate from 60 to 100 days of gestation (DG) and from 2 to 14 days post partum (DP). This analysis showed that lymphoid aggregations first appear at DG65, gradually develop during the fetal stage, and expand after birth. In addition, the mRNA expression of chemokine genes involved in lymphoid aggregation and localization was analyzed. CCL19 expression showed the most marked increase and a sharp peak after birth. CCL21 expression changed moderately but showed an interesting bimodal pattern. CXCL13 expression steadily increased throughout the study period. Thus, we demonstrated the mRNA expression of chemokine characteristically changed accompanying tonsillar development.

著者

Naito Hiroyuki Nakamura Amane Inoue Maki

出版者

公益社団法人 日本実験動物学会

雑誌

Experimental animals (ISSN:13411357)

巻号頁・発行日

vol.47, no.2, pp.89-96, 1998-04

参考文献数

39

被引用文献数

4 11

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The ontogenetic changes in responsiveness to benzodiazepine receptor ligands on ultrasonic vocalizations in rat pups from the age of day 3 to day 12 were evaluated. Rat pups, while separated from their dam and littermates and placed in a cold environment, emit ultrasonic vocalizations. These ultrasonic calls became attenuated dose-dependently in number and power after administration of the anxiolytic diazepam (0.25-1.0 mg/kg, sc), but the inhibitory effect of diazepam at the highest dose was less on day 6 and day 9. Moreover, type 1 benzodiazepine receptor ligands, Ro16-6028 and Ro23-0364 (0.5-2.0 mg/kg, sc), also dose-dependently attenuated the ultrasonic vocalizations 30-60 min after injection. The inhibitory effects of these drugs became more pronounced with the increasing age of the pup, and they were equivalent on day 12 to those in adult rats. These results suggest that different ontogenetic changes in development of two subtypes of central benzodiazepine receptors of pups might be related in the psychopharmacological mediation of the ultrasonic vocalization.<br>

著者

Abdulsamad Alsalahi Mohammed A. Alshawsh Zamri Chik Zahurin Mohamed

出版者

Japanese Association for Laboratory Animal Science

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

vol.67, no.4, pp.517-526, 2018 (Released:2018-11-01)

参考文献数

42

被引用文献数

9

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People consume Catha edulis (khat) for its euphoric effect, and type 1 diabetics have claimed that khat could reduce elevated levels of blood sugar. However, khat has been suggested to provoke diabetes mellitus through destruction of pancreatic β-cells. This study investigated the effect of an ethanolic khat extract on pancreatic functions in type 1 diabetes (T1DM)-induced male Sprague-Dawley rats and to assess its in vitro cytotoxicity in rat pancreatic β-cells (RIN-14B). T1DM was induced in a total of 20 rats with a single intraperitoneal injection of 75 mg/kg of streptozotocin. The rats were distributed into four groups (n=5): the diabetic control, 8 IU insulin-treated, 200 mg/kg khat-treated, and 400 mg/kg khat-treated groups. Another 5 rats were included as a nondiabetic control. Body weight, fasting blood sugar, and caloric intake were recorded weekly. Four weeks after treatment, the rats were sacrificed, and blood was collected for insulin, lipid profile, total protein, amylase, and lipase analysis, while pancreases were harvested for histopathology. In vitro, khat exerted moderate cytotoxicity against RIN-14B cells after 24 and 48 h but demonstrated greater inhibition against RIN-14B cells after 72 h. Neither 200 mg/kg nor 400 mg/kg of khat produced any significant reduction in blood sugar; however, 200 mg/kg khat extract provoked more destruction of pancreatic β-cells as compared with the diabetic control. Ultimately, neither 200 mg/kg nor 400 mg/kg of khat extract could produce a hypoglycemic effect in T1DM-induced rats. However, 200 mg/kg of khat caused greater destruction of pancreatic β-cells, implying that khat may cause a direct cytotoxic effect on pancreatic β-cells in vitro.

著者

Jun IWAMOTO Azusa SEKI Tsuyoshi TAKEDA Yoshihiro SATO Harumoto YAMADA James K. YEH

出版者

Japanese Association for Laboratory Animal Science

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

vol.67, no.3, pp.R1, 2018 (Released:2018-07-30)

被引用文献数

1

---

The following articles have been retracted by the Editorial Board of Experimental Animals, because some parts of their contents were published elsewhere.

著者

Jun IWAMOTO Azusa SEKI Tsuyoshi TAKEDA Yoshihiro SATO Harumoto YAMADA James K. YEH

出版者

Japanese Association for Laboratory Animal Science

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

vol.67, no.3, pp.R2, 2018 (Released:2018-07-30)

被引用文献数

1

---

The following articles have been retracted by the Editorial Board of Experimental Animals, because some parts of their contents were published elsewhere.

著者

Kei MATSUNO Setsunosuke IHARA

出版者

公益社団法人 日本実験動物学会

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

vol.64, no.3, pp.323-332, 2015 (Released:2015-08-01)

参考文献数

16

被引用文献数

3

著者

Yunyun TIAN Li TAO Siriguleng ZHAO Dapeng TAI Dongjun LIU Pengxia LIU

出版者

公益社団法人 日本実験動物学会

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

pp.15-0031, (Released:2015-11-27)

被引用文献数

15

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Mesenchymal stem cells (MSCs) are one of the most promising cell populations for tissue engineering and regenerative medicine. Of utmost importance to MSC research is identification of MSC sources that are easily obtainable and stable. Several studies have shown that MSCs can be isolated from amniotic fluid. The sheep is one of the main types of farm animal, and it has many biophysical and biochemical similarities to humans. Here, we obtained MSCs from ovine amniotic fluid and determined the expansion capacity, surface and intracellular marker expression, karyotype, and multilineage differentiation ability of these ovine amniotic fluid mesenchymal stem cells (oAF-MSCs). Moreover, expression levels of differentiation markers were measured using reverse transcription-qPCR (RT-qPCR). Our phenotypic analysis shows that the isolated oAF-MSCs are indeed MSCs.

著者

目加田 和之 阿部 訓也 村上 亜弓 NAKAMURA Satoe NAKATA Hatsumi MORIWAKI Kazuo OBATA Yuichi YOSHIKI Atsushi

出版者

Japanese Association for Laboratory Animal Science

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

vol.58, no.2, pp.141-149, 2009

被引用文献数

1 267

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The C57BL/6 mouse is the most well-known inbred mouse strain, and has been widely used as a genetic background for congenic and mutant mice. A number of C57BL/6 substrains have been derived from the C57BL/6 founder line and are reported to differ in several phenotypes. There are several major sources of C57BL/6 substrains for the biomedical research community. The importance of their genetic and phenotypic differences among substrains, however, has not yet been well recognized by biomedical researchers. Here, we report the result of screening of the functional deletion of the nicotinamide nucleotide transhydrogenase (<i>Nnt</i>) gene and 1,446 SNPs genotyping among seven C57BL/6 substrains from different sources, such as C57BL/6J, C57BL/6JJcl, C57BL/6JJmsSlc, C57BL/6NJcl, C57BL/6NCrlCrlj, C57BL/6NTac, and C57BL/6CrSlc. The deletion of exon 7-11 in the <i>Nnt</i> gene that was previously reported in C57BL/6J was also observed in other C57BL/6J substrains, indicating that this functional deletion probably occurred at an early stage in the establishment of C57BL/6J substrains. The genotyping of SNP loci clearly demonstrate genetic differences between C57BL/6J and C57BL/6N substrains at 11 loci. Besides, we found another SNP differing between C57BL/6J and other C57BL/6J substrains available from commercial breeders. No genetic difference was detected among C57BL/6N substrains. The C57BL/6CrSlc mouse, originally derived from the National Cancer Institute of the NIH was found to be the same as the C57BL/6N substrains by the SNP pattern. These data will be useful for accurate genetic monitoring of genetically engineered mice with the C57BL/6 background.<br>

著者

An-Yun GUO Kwok-Sui LEUNG Ming-Fai SIU Jiang-Hui QIN Simon Kwoon-Ho CHOW Ling QIN Chi-yu LI Wing-Hoi CHEUNG

出版者

公益社団法人 日本実験動物学会

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

pp.15-0025, (Released:2015-07-17)

被引用文献数

2 46

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Sarcopenia is an age-related systemic syndrome with progressive deterioration in skeletal muscle functions and loss in mass. Although the senescence-accelerated mouse P8 (SAMP8) was reported valid for muscular ageing research, there was no report on the details such as sarcopenia onset time. Therefore, this study was to investigate the change of muscle mass, structure and functions during the development of sarcopenia. Besides the average life span, muscle mass, structural and functional measurements were also studied. Male SAMP8 animals were examined at month 6, 7, 8, 9, and 10, in which the right gastrocnemius was isolated and tested for ex vivo contractile properties and fatigability while the contralateral one was harvested for muscle fiber cross-sectional area (FCSA) and typing assessments. Results showed that the peak of muscle mass appeared at month 7 and the onset of contractility decline was observed from month 8. Compared with month 8, most of the functional parameters at month 10 decreased significantly. Structurally, muscle fiber type IIA made up the largest proportion of the gastrocnemius, and the fiber size was found to peak at month 8. Based on the altered muscle mass, structural and functional outcomes, it was concluded that the onset of sarcopenia in SAMP8 animals was at month 8. SAMP8 animals at month 8 should be at pre-sarcopenia stage while month 10 at sarcopenia stage. It is confirmed that SAMP8 mouse can be used in sarcopenia research with established time line in this study.

著者

Kenji SAKUMA Susumu HAYASHI Yoshiyuki YASAKA Hiroto NISHIJIMA Hisakage FUNABASHI Masayoshi HAYASHI Hideaki MATSUOKA Mikako SAITO

出版者

公益社団法人 日本実験動物学会

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

vol.62, no.2, pp.101-107, 2013 (Released:2013-04-25)

参考文献数

16

被引用文献数

4

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In order to provide healthy experimental animals, it is important to find and remove animals that have been accidentally exposed to various stresses during breeding. This study focuses mouse health-care management. Here we used human olfaction and gas chromatography-mass spectrometry to assess odor intensity and determine the concentrations of odor components. The feces were collected from mice that were exposed to 4 different stresses (no bedding chips, shaking, fasting, and movement restriction). These stresses caused a change in odor intensity as assessed by 6 panelists. Seventeen components were identified as dominant components in the odor that was emitted from feces. The concentration of each compound was converted to relative values versus its odor threshold levels in order to select ones effective for the quality of the odor. As a result, 12 selected components were found to be a useful set for the recognition of mice bred under different stress conditions. The present results may provide useful information for the development of standard fecal odor materials that may be used for the training of mouse care personnel.

著者

Francesco FAZIO Stefania CASELLA Claudia GIANNETTO Elisabetta GIUDICE Giuseppe PICCIONE

出版者

公益社団法人 日本実験動物学会

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

pp.14-0032, (Released:2014-08-05)

被引用文献数

2 18

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Haptoglobin (Hp), serum amyloid A (SAA), C-reactive protein (CRP), white blood cells (WBC), reactive oxygen metabolites (ROMs), the antioxidant barrier (Oxy-adsorbent) and thiol groups of plasma compounds (SHp) were measured in ten dogs that had been transported a distance of about 230 km within 2 h (experimental group) and in ten dogs that had not been subjected to road transportation (control group). Blood was collected via cephalic venipuncture before road transportation (T0), after road transportation (T1), and more than 6 (T6) and 24 (T24) hours after road transportation in the experimental group (Group A) and at the same time points in the control group (Group B). The GLM (general linear model) Repeated Measures procedure showed a significant difference between the two groups (P<0.0001) and a significant rise (P<0.0001) in the concentrations of Hp, SAA, CRP, WBC, ROMs, Oxy-adsorbent and SHp after road transportation in Group A, underlining that physiological and homeostatic mechanisms are modified differently at various sampling times.

著者

Ruixia Zeng YiBo Zhang Peng Du

出版者

公益社団法人 日本実験動物学会

雑誌

Experimental Animals (ISSN:13411357)

巻号頁・発行日

vol.63, no.1, pp.73-78, 2014 (Released:2014-02-07)

参考文献数

19

被引用文献数

3 18

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Melanocortin 4 receptor (MC4R), which is associated with inherited human obesity, is involoved in food intake and body weight of mammals. To study the relationships between MC4R gene polymorphism and body weight in Beagle dogs, we detected and compared the nucleotide sequence of the whole coding region and 3′- and 5′- flanking regions of the dog MC4R gene (1214 bp). In 120 Beagle dogs, two SNPs (A420C, C895T) were identified and their relation with body weight was analyzed with RFLP-PCR method. The results showed that the SNP at A420C was significantly associated with canine body weight trait when it changed amino acid 101 of the MC4R protein from asparagine to threonine, while canine body weight variations were significant in female dogs when MC4R nonsense mutation at C895T. It suggested that the two SNPs might affect the MC4R gene’s function which was relative to body weight in Beagle dogs. Therefore, MC4R was a candidate gene for selecting different size dogs with the MC4R SNPs (A420C, C895T) being potentially valuable as a genetic marker.