Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (19): 4400-4416.doi: 10.3864/j.issn.0578-1752.2026.19.017

• ANIMAL SCIENCE·VETERINARY SCIENCE • Previous Articles     Next Articles

Differential Analysis of Hypoxia-Related Factors in the Small Intestines of Yaks and Yellow Cattle

ZHANG YeQiong1(), WANG Jun1, XU MingWei1, HU JiaQi1, LIU ShaSha1, LAN WenLin1, TAN Xiao1, YANG Kun1,2,3()   

  1. 1 College of Life Science and Engineering, Northwest Minzu University, Lanzhou 730000
    2 Gansu Provincial Technology Innovation Center for Animal Cells, Biomedical Research Center, Northwest Minzu University, Lanzhou 730030
    3 Key Laboratory of Bioengineering and Technology, State Ethnic Affairs Commission, Biomedical Research Center, Northwest Minzu University, Lanzhou 730030
  • Received:2025-12-29 Accepted:2026-04-10 Online:2026-10-01 Published:2026-10-10
  • Contact: YANG Kun

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Abstract:

【Objective】This study aimed to investigate the differential expression of hypoxia-related factors—Pyruvate dehydrogenase kinase 1 (PDK1), Hypoxia-inducible factor-1α (HIF-1α), Transforming growth factor-β (TGF-β), Vascular endothelial growth factor (VEGF), Mothers against decapentaplegic homolog 2 (Smad2), and Mothers against decapentaplegic homolog 3 (Smad3)—in the small intestinal tissues of yaks, to elucidate the intestinal adaptation mechanisms of yaks to hypoxic environments.【Method】Small intestinal tissues (duodenum, jejunum, and ileum) from plateau yaks and plain yellow cattle were selected as research subjects. Hematoxylin-eosin staining (H.E), periodic acid-Schiff staining (PAS), Masson’s trichrome staining, and immunohistochemical staining were performed. Tissue sections after staining were photographed under a light microscope. Image analysis software was used to measure villus height (V) and crypt depth (C) in H.E-stained sections, as well as the average optical density (AOD) of PAS, Masson, and immunohistochemical staining. The average optical density value was employed to represent positive expression results, thereby investigating the distribution characteristics and expression differences of hypoxia-related factors in different species and intestinal segments.【Result】H.E staining results indicated that the V/C ratio (villus height/crypt depth), an indicator of intestinal digestion, in both yaks and yellow cattle was significantly higher in the duodenum compared to the jejunum and ileum (P<0.05). Furthermore, the duodenal V/C ratio of yaks was significantly higher than that of yellow cattle (P<0.05). PAS staining showed that PAS-positive substances were expressed with significantly higher intensity in the duodenum than in the jejunum and ileum (P<0.05), with strong positive expression observed in the striated border, goblet cells, and Paneth cells. The expression in all intestinal segments of yaks was significantly higher than in yellow cattle (P<0.05). Masson's trichrome staining results revealed significant interspecies and segmental differences in collagen fiber content within the small intestinal tissues of both yaks and yellow cattle (P<0.05). Specifically, the proportion of collagen fibers in the duodenum and jejunum of yellow cattle was significantly higher than in the corresponding segments of yaks (P<0.05). Immunohistochemical analysis detected distinct positive brown-yellow expression of PDK1, HIF-1α, TGF-β, and VEGF in the small intestinal tissues, primarily localized to the striated border, Paneth cells, and goblet cells. Differences in intestinal gland expression were observed between species: weak positive expression was seen in the duodenal and jejunal glands of yellow cattle, while strong positive expression was observed in the corresponding locations in yaks. Tissue segment analysis showed that the expression levels of these factors were significantly higher in the duodenum than in the jejunum (P<0.05). Regarding species differences, the overall expression levels in yak small intestinal tissues were significantly higher than in yellow cattle (P<0.05). However, the expression patterns of Smad2 and Smad3 differed from those of the aforementioned factors. The expressions of Smad2 and Smad3 in the duodenum were significantly lower than those in the ileum and jejunum, respectively (P<0.05). Furthermore, the expression of Smad2/3 in the duodenum and ileum of yaks was significantly lower than in yellow cattle (P<0.05), while in the jejunum of yaks, it was significantly higher than in yellow cattle (P<0.05).【Conclusion】Results from H&E staining, special staining, and immunohistochemical staining revealed significant differences between yaks and yellow cattle in the morphological structure, carbohydrate distribution, and expression of hypoxia-related factors in the small intestine. These findings suggest that yak intestines possess unique structural and molecular adaptive features to hypoxia, and that factors such as PDK1, HIF-1α, and VEGF may play important roles in the intestinal hypoxic adaptation mechanisms of yaks. This study provides fundamental data for research on intestinal adaptation in high-altitude animals and offers a reference for exploring the mechanisms and prevention of hypoxia-related intestinal diseases.

Key words: yak, small intestine, hypoxia, pyruvate dehydrogenase kinase 1 (PDK1), hypoxia inducible factor 1α (HIF-1α), vascular endothelial growth factor (VEGF)

Fig. 1

Small intestinal tissue sections of yak and yellow cattle (H.E staining) A-C, J-L are H.E staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 200×; D-F and M-O are the H.E staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 400×; G-I and P-R are H.E staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 1000×. (IG: Intestinal gland DG: Duodenal gland. GC: Goblet cells. ML: Muscle layer. AC: Absorptive cells. PC: Paneth cells. MM: Muscularis mucosae. BV: Blood vessels. SB: Striated border. S: Submucosa. The same as below"

Fig. 2

Morphological measurement of small intestinal tissue in Yaks and yellow cattle (H.E staining) (Ⅰ) Comparison of intestinal villus height among different tissues and species; (Ⅱ) Comparison of crypt depth among different tissues and species; (Ⅲ) Comparison of digestive index (V/C ratio) among different tissues and species. Wherein, ns indicates no significant difference between species (P > 0.05), * indicates a significant difference between species (P < 0.05), ** indicates a very significant difference between species (P < 0.01), and *** indicates an extremely significant difference between species (P < 0.001); different uppercase letters (A, B, C) indicate significant differences among different small intestinal tissues of yellow cattle (P < 0.05), and different lowercase letters (a, b, c) indicate significant differences among different small intestinal tissues of yaks (P < 0.05). The same as below"

Fig. 3

Small intestinal tissue sections of yak and yellow cattle (PAS staining) A-C, J-L are PAS staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 200×; D-F and M-O are the PAS staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 400×; G-I and P-R are PAS staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 1000×"

Fig. 4

Morphological measurement of small intestinal tissue in yaks and yellow cattle (PAS staining)"

Fig. 5

Small intestinal tissue sections of yak and yellow cattle (Masson staining) A-C, J-L are Masson staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 200×; D-F and M-O are the Masson staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 400×; G-I and P-R are Masson staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 1000×"

Fig. 6

Morphological measurement of small intestinal tissue in yaks and yellow cattle (Masson staining)"

Fig. 7

Small intestinal tissue sections of yak and yellow cattle (negative control) A-C, J-L are the negative control group of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 200×; D-F and M-O are the negative control group of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 400×; G-I and P-R are the negative control group staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 1000×"

Fig. 8

Small intestinal tissue sections of yak and yellow cattle (PDK1 staining) A-C, J-L are PDK1 staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 200×; D-F and M-O are PDK1 staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 400×; G-I and P-R are PDK1 staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 1000×"

Fig. 9

Morphological measurement of small intestinal tissue in Yaks and yellow cattle (PDK1 staining)"

Fig. 10

Small intestinal tissue sections of yak and yellow cattle (HIF-1α staining) A-C, J-L are HIF-1α staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 200×; D-F and M-O are the HIF-1α staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 400×; G-I and P-R are HIF-1α staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 1000×"

Fig. 11

Morphological measurement of small intestinal tissue in Yaks and yellow cattle (HIF-1α staining)"

Fig. 12

Small intestinal tissue sections of yak and yellow cattle (VEGF staining) A-C, J-L are VEGF staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 200×; D-F and M-O were the VEGF staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 400×; G-I and P-R were VEGF staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 1000×"

Fig. 13

Morphological measurement of small intestinal tissue in Yaks and yellow cattle (VEGF staining)"

Fig. 14

Small intestinal tissue sections of yak and yellow cattle (TGF-β staining) A-C, J-L are TGF-β staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 200×; D-F and M-O are the TGF-β staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 400×; G-I and P-R are TGF-β staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 1000×"

Fig. 15

Morphological measurement of small intestinal tissue in Yaks and yellow cattle (TGF-β staining)"

Fig. 16

Small intestinal tissue sections of yak and yellow cattle (Smad2 staining) A-C, J-L are Smad2 staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 200×; D-F and M-O are the Smad2 staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 400×; G-I and P-R are Smad2 staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 1000×"

Fig. 17

Morphological measurement of small intestinal tissue in Yaks and yellow cattle (Smad2 staining)"

Fig. 18

Small intestinal tissue sections of yak and yellow cattle (Smad3 staining) A-C, J-L are Smad3 staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 200×; D-F and M-O are the Smad3 staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 400×; G-I and P-R are Smad3 staining of the duodenum, jejunum and ileum of yellow cattle and yaks in sequence, 1000×"

Fig. 19

Morphological measurement of small intestinal tissue in Yaks and yellow cattle (Smad3 staining)"

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