食物单宁和蛋白对小家鼠（Mus musculus）的母体效应研究
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摘要
母体效应是指双亲除基因遗传之外的其它方式对其后代表型的影响,该效应对后代形态、生理及行为影响很大且持续终生。它是子代对环境异质性的一种表型反应,是种群进化的一种动力,也可能是种群调节的一种机制。本论文以活捕于河南新郑地区的野生小家鼠(Mus musculus)及其室内繁殖后代为研究对象,采用正、负营养因子双因素协同作用的方法,设置食物单宁酸浓度为0%和5%,蛋白质水平为15%、20%和25%,人工配制成6种饲料。向处于妊娠期和非繁殖期的成年健康个体分别投喂不同配制饲料2 W,测定小家鼠的蛋白质代谢、消化道形态、免疫、繁殖和社群行为等指标,探讨母体妊娠期营养对子代生长发育、消化道形态、免疫、繁殖及社群、筑巢行为的效应。本文还测定了年龄对小家鼠筑巢的影响。
主要研究结果和结论如下：
1)食物中添加单宁和蛋白对成年小家鼠摄食量均无显著影响,但与24 h粪便量及粪样N含量均呈显著正相关关系；食物添加单宁对小鼠体重具有显著抑制作用,而食物添加蛋白对小鼠体重作用不显著。结果表明,食物添加单宁对成年小家鼠蛋白质消化率和体重均有显著的抑制作用,食物中添加蛋白质的影响不明显。
2)食物添加单宁对成年小家鼠小肠、结肠干重和小肠长度等具有显著抑制作用,食物蛋白水平则与小肠干重等呈正相关关系,而胃肠重量及长度与食物质量相关性不显著。在妊娠期投喂配制饲料小家鼠的消化系统重量明显高于非繁殖期。结果表明,动物消化道形态具有一定可塑性,能根据能量需求程度与食物质量进行相应调整。
3)食物中添加单宁可显著增加小家鼠肾上腺指数及粪样皮质激素水平,对脾脏指数作用不显著；而食物添加蛋白对脾脏指数具有显著增加效应,对肾上腺指数及粪样皮质激素作用不显著；小家鼠妊娠期脾脏指数显著增高,而肾上腺指数及粪样皮质激素显著低于非繁殖个体。子宫指数、睾酮、雌二醇含量等均与食物质量和动物能量需求程度呈显著正相关关系。
4)食物添加单宁对雌性与雄性间攻击行为(FMA)频次具有一定的抑制作用,雄性与雄性间防御行为(MMD)频次与食物蛋白水平负相关,其它的性别组合间攻击、防御、社会及非社会行为频次与食物质量均无显著相关性,且动物攻击、防御行为频次均处于较低水平；在妊娠期,同性或异性小鼠间攻击、防御行为频次与非繁殖期间攻击、防御行为频次间无显著性差异,而非社会行为频次明显高于非繁殖期间的非社会行为频次,同时社会行为频次低于非生育组。这可能是小家鼠通过减少活动以降低能量支出的一种行为调节机制。
5)小家鼠母体妊娠期投喂配制饲料各组间子代性比无显著差异,组间体重、存活率在断奶后差异性存在增大趋势,且与母体妊娠期食物质量呈正相关关系,但均未达到显著性差异水平；母体妊娠期食物添加单宁或蛋白含量低,对子代形态发育、神经发育均具有显著的抑制作用；母体妊娠期食物添加单宁对平甲状腺激素含量均无显著影响,而母体妊娠期食物高蛋白组胎仔数显著高于中蛋白组胎仔数,其它蛋白组间胎仔数差异性均不显著,妊娠期食物蛋白水平与血清三碘甲状腺氨酸含量相一致,与甲状腺素含量相反。
6)小家鼠母体妊娠期食物添加单宁对子代胃干重、结肠干重、小肠长度等具有显著的抑制作用,盲肠长度与母体食物蛋白质水平正相关；母体妊娠期食物添加单宁能增加子代肾上腺重量及皮质酮含量,抑制睾丸重量,而脾脏重量和粪样睾酮含量与蛋白质水平正相关；结果表明,母体妊娠期食物营养胁迫能通过母体效应对子代消化道形态、免疫及繁殖功能产生一定影响。
7)小家鼠母体妊娠期食物添加单宁对子代雄性的筑巢能力具有显著抑制作用,而母体妊娠期食物蛋白质水平与雌性的筑巢能力呈一定正相关关系；母体妊娠期食物单宁和蛋白对子代社群行为影响均不明显。
8)小家鼠在亚成年期已具备较强的筑巢能力,成年与亚成年小家鼠能够快速完成筑巢,24 h即可筑成稳定的杯状巢,而幼年组未能筑成稳定巢,小家鼠雄性与雌性筑巢能力相当,连续4d巢材获取重量表现为成年组>亚成年组,而4d后两组间筑巢等级无显著差异,这与两组间身体大小有关。
综上所述,食物单宁和蛋白对小家鼠蛋白质消化率、消化道形态、免疫、繁殖功能和社群行为具有重要作用,并能通过母体效应对子代形态发育、神经发育、消化道形态、免疫、繁殖功能及筑巢行为产生重要影响。食物单宁和蛋白胁迫引起的母体效应可能是小家鼠种群动态的重要调节因素之一。
Maternal effect is a situation where the phenotype of an organism is determined not only by the environment it experiences and its genotype, but also by the environment and phenotype of its mother, and can be found in several aspects including morphology, physiology and behavior of offsprings. This effect is also a phenotype reaction of offsprings themselves to the environmental heterogeneity. It is an important source of evolutionary dynamics, and might be a mechanism of population regulation. In this research, the maternal effect of food quality on house mice (Mus musculus) was studied, and we aimed to understand how the nutritious conditions during gestation of parents influence its offspring. The experimental food with different tannin (0% and 5%) and protein (15%,20% and 25%) were provided to healthy adult breeding groups and nonbreeding groups respectively for two weeks. Then we studied the effect of food with different tannic and protein on protein metabolistic rate, digestive tract morphology, immunological status, reproductive and agonistic behaviors, maternal nutrition on growing development, digestive tract morphology, immunology, reproductive and agonistic, nesting behavior of offspring in house mice (Mus musculus). This paper also tested the effects of age on nest-building of house mice.
The major results from this research are as follows:
1) Food with different tannin or protein had no obvious effect on food-intake in mice, however, significant positive correlation between feces amount in 24 hours and protein content. Tannin-added food would inhibit the increase of bodymass of adult non-breeding individuals, but there is no significant difference among protein groups. This indicated that tannin-based food can inhibit the increase of bodyweight and impact protein digestibility.
2) Fed with tannin-added food have a significant inhibition on the dry weight of small intestine or colon and the length of small intestine of house mice; while the protein content positively correlated to dry weight of small intestine; but there is no significant difference between the other length and mass of digestive tracts and food quality. The weight of digestive system during pregnancy is heavier than that in nonbreeding period. Animal can adjust their digestive tract to certain extent based on trade-off between energy need and food quality.
3) Fed with tannin-added food can greatly increase adrenal gland index and cortin content, but there is no significant inhibition on the index of spleen; however, the protein content positively correlated to index of spleen, but there is no significant difference with adrenal gland index and cortin content; the index of spleen were greatly increase during breeding period. The uterine index, testosterone, estradiol levels all display a significant positive correlation with food quality and demand of energy.
4) Tannin-added food would partly restrain aggressive behavior between female and male (FMA), while defending behavior between male and male (MMD) exhibited a negative relationship with protein level in food, but there is no significant difference between other agonistic behaviors frequence and food quality and the attacking behavior and the defensive behavior show a low level. The breeding has a positive relationhip with non-social behavior, but a negative relationship for social behavior. This may be a regulating mechanism of house mice to decrease the energy expenditure by reducing activity.
5) During pregnant period fed with food of different tannin and protein have no significant influences on sex ratio, bodymass, and survival of offspring, but can greatly inhibit morphological and neurological development of offspring. The protein levels positively correlated to litter size and triiodothyronine content in serum, but a negative relationship for Thyroxine content.
6) During pregnant period, tannin-added food has a significant inhibition on the dry weight of stomach, colon and length of small intestine, and testicular index; ehereas increase the weight of adrenal gland and the corticosterone levels in dejecta. However, the length of caecum and testosterone levels were both positively correlated with protein levels. This implied that deficient food nutrition during pregnancy would affect digestive tract traits, immunology and reproductive ability of offspring through maternal effect.
7) Pregnant females fed tannin-added food would prohibit nesting ability of male offsprings; while the nesting ability of female offsprings was weakly and positively related to food proteinlevels. Food with different tannin and protein levels in during pregnant perriod exerted insignificant influence on agonistic behavior.
8) House mice behaved good nesting capacity during sub-adulthood, the house mice nested well in both adult and subadult stages, because they could complete a stable cup-shaped nest, while the juvenile group could not establish stable nests, the difference of nesting capacity between male and female was not significant; and although the amount of cotton, nesting material, used by adults in four days were significantly higher than that in sub-adults, the nesting degree between the two groups was not significantly different after four days, which was related to the body size of experimental animals.
In summary, the tannin and protein levels in food played important roles on protein digestibility, gastrointestinal morphology, immune, reproductive ability and social behavior, and showed great effects on morphological and neural development, digestive tract morphology, immunology, reproductive function and nesting behavior via maternal effects. We thus claimed that tannin and protein in food may be one of important regulators of population dynamics.
主要研究结果和结论如下：
1)食物中添加单宁和蛋白对成年小家鼠摄食量均无显著影响,但与24 h粪便量及粪样N含量均呈显著正相关关系；食物添加单宁对小鼠体重具有显著抑制作用,而食物添加蛋白对小鼠体重作用不显著。结果表明,食物添加单宁对成年小家鼠蛋白质消化率和体重均有显著的抑制作用,食物中添加蛋白质的影响不明显。
2)食物添加单宁对成年小家鼠小肠、结肠干重和小肠长度等具有显著抑制作用,食物蛋白水平则与小肠干重等呈正相关关系,而胃肠重量及长度与食物质量相关性不显著。在妊娠期投喂配制饲料小家鼠的消化系统重量明显高于非繁殖期。结果表明,动物消化道形态具有一定可塑性,能根据能量需求程度与食物质量进行相应调整。
3)食物中添加单宁可显著增加小家鼠肾上腺指数及粪样皮质激素水平,对脾脏指数作用不显著；而食物添加蛋白对脾脏指数具有显著增加效应,对肾上腺指数及粪样皮质激素作用不显著；小家鼠妊娠期脾脏指数显著增高,而肾上腺指数及粪样皮质激素显著低于非繁殖个体。子宫指数、睾酮、雌二醇含量等均与食物质量和动物能量需求程度呈显著正相关关系。
4)食物添加单宁对雌性与雄性间攻击行为(FMA)频次具有一定的抑制作用,雄性与雄性间防御行为(MMD)频次与食物蛋白水平负相关,其它的性别组合间攻击、防御、社会及非社会行为频次与食物质量均无显著相关性,且动物攻击、防御行为频次均处于较低水平；在妊娠期,同性或异性小鼠间攻击、防御行为频次与非繁殖期间攻击、防御行为频次间无显著性差异,而非社会行为频次明显高于非繁殖期间的非社会行为频次,同时社会行为频次低于非生育组。这可能是小家鼠通过减少活动以降低能量支出的一种行为调节机制。
5)小家鼠母体妊娠期投喂配制饲料各组间子代性比无显著差异,组间体重、存活率在断奶后差异性存在增大趋势,且与母体妊娠期食物质量呈正相关关系,但均未达到显著性差异水平；母体妊娠期食物添加单宁或蛋白含量低,对子代形态发育、神经发育均具有显著的抑制作用；母体妊娠期食物添加单宁对平甲状腺激素含量均无显著影响,而母体妊娠期食物高蛋白组胎仔数显著高于中蛋白组胎仔数,其它蛋白组间胎仔数差异性均不显著,妊娠期食物蛋白水平与血清三碘甲状腺氨酸含量相一致,与甲状腺素含量相反。
6)小家鼠母体妊娠期食物添加单宁对子代胃干重、结肠干重、小肠长度等具有显著的抑制作用,盲肠长度与母体食物蛋白质水平正相关；母体妊娠期食物添加单宁能增加子代肾上腺重量及皮质酮含量,抑制睾丸重量,而脾脏重量和粪样睾酮含量与蛋白质水平正相关；结果表明,母体妊娠期食物营养胁迫能通过母体效应对子代消化道形态、免疫及繁殖功能产生一定影响。
7)小家鼠母体妊娠期食物添加单宁对子代雄性的筑巢能力具有显著抑制作用,而母体妊娠期食物蛋白质水平与雌性的筑巢能力呈一定正相关关系；母体妊娠期食物单宁和蛋白对子代社群行为影响均不明显。
8)小家鼠在亚成年期已具备较强的筑巢能力,成年与亚成年小家鼠能够快速完成筑巢,24 h即可筑成稳定的杯状巢,而幼年组未能筑成稳定巢,小家鼠雄性与雌性筑巢能力相当,连续4d巢材获取重量表现为成年组>亚成年组,而4d后两组间筑巢等级无显著差异,这与两组间身体大小有关。
综上所述,食物单宁和蛋白对小家鼠蛋白质消化率、消化道形态、免疫、繁殖功能和社群行为具有重要作用,并能通过母体效应对子代形态发育、神经发育、消化道形态、免疫、繁殖功能及筑巢行为产生重要影响。食物单宁和蛋白胁迫引起的母体效应可能是小家鼠种群动态的重要调节因素之一。
Maternal effect is a situation where the phenotype of an organism is determined not only by the environment it experiences and its genotype, but also by the environment and phenotype of its mother, and can be found in several aspects including morphology, physiology and behavior of offsprings. This effect is also a phenotype reaction of offsprings themselves to the environmental heterogeneity. It is an important source of evolutionary dynamics, and might be a mechanism of population regulation. In this research, the maternal effect of food quality on house mice (Mus musculus) was studied, and we aimed to understand how the nutritious conditions during gestation of parents influence its offspring. The experimental food with different tannin (0% and 5%) and protein (15%,20% and 25%) were provided to healthy adult breeding groups and nonbreeding groups respectively for two weeks. Then we studied the effect of food with different tannic and protein on protein metabolistic rate, digestive tract morphology, immunological status, reproductive and agonistic behaviors, maternal nutrition on growing development, digestive tract morphology, immunology, reproductive and agonistic, nesting behavior of offspring in house mice (Mus musculus). This paper also tested the effects of age on nest-building of house mice.
The major results from this research are as follows:
1) Food with different tannin or protein had no obvious effect on food-intake in mice, however, significant positive correlation between feces amount in 24 hours and protein content. Tannin-added food would inhibit the increase of bodymass of adult non-breeding individuals, but there is no significant difference among protein groups. This indicated that tannin-based food can inhibit the increase of bodyweight and impact protein digestibility.
2) Fed with tannin-added food have a significant inhibition on the dry weight of small intestine or colon and the length of small intestine of house mice; while the protein content positively correlated to dry weight of small intestine; but there is no significant difference between the other length and mass of digestive tracts and food quality. The weight of digestive system during pregnancy is heavier than that in nonbreeding period. Animal can adjust their digestive tract to certain extent based on trade-off between energy need and food quality.
3) Fed with tannin-added food can greatly increase adrenal gland index and cortin content, but there is no significant inhibition on the index of spleen; however, the protein content positively correlated to index of spleen, but there is no significant difference with adrenal gland index and cortin content; the index of spleen were greatly increase during breeding period. The uterine index, testosterone, estradiol levels all display a significant positive correlation with food quality and demand of energy.
4) Tannin-added food would partly restrain aggressive behavior between female and male (FMA), while defending behavior between male and male (MMD) exhibited a negative relationship with protein level in food, but there is no significant difference between other agonistic behaviors frequence and food quality and the attacking behavior and the defensive behavior show a low level. The breeding has a positive relationhip with non-social behavior, but a negative relationship for social behavior. This may be a regulating mechanism of house mice to decrease the energy expenditure by reducing activity.
5) During pregnant period fed with food of different tannin and protein have no significant influences on sex ratio, bodymass, and survival of offspring, but can greatly inhibit morphological and neurological development of offspring. The protein levels positively correlated to litter size and triiodothyronine content in serum, but a negative relationship for Thyroxine content.
6) During pregnant period, tannin-added food has a significant inhibition on the dry weight of stomach, colon and length of small intestine, and testicular index; ehereas increase the weight of adrenal gland and the corticosterone levels in dejecta. However, the length of caecum and testosterone levels were both positively correlated with protein levels. This implied that deficient food nutrition during pregnancy would affect digestive tract traits, immunology and reproductive ability of offspring through maternal effect.
7) Pregnant females fed tannin-added food would prohibit nesting ability of male offsprings; while the nesting ability of female offsprings was weakly and positively related to food proteinlevels. Food with different tannin and protein levels in during pregnant perriod exerted insignificant influence on agonistic behavior.
8) House mice behaved good nesting capacity during sub-adulthood, the house mice nested well in both adult and subadult stages, because they could complete a stable cup-shaped nest, while the juvenile group could not establish stable nests, the difference of nesting capacity between male and female was not significant; and although the amount of cotton, nesting material, used by adults in four days were significantly higher than that in sub-adults, the nesting degree between the two groups was not significantly different after four days, which was related to the body size of experimental animals.
In summary, the tannin and protein levels in food played important roles on protein digestibility, gastrointestinal morphology, immune, reproductive ability and social behavior, and showed great effects on morphological and neural development, digestive tract morphology, immunology, reproductive function and nesting behavior via maternal effects. We thus claimed that tannin and protein in food may be one of important regulators of population dynamics.
引文
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