棕色田鼠社会识别行为的研究
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摘要
化学信号深刻地影响着啮齿动物的行为，并在其生存中有重要意义。啮齿动物的社会关系依赖于社会识别，这种识别包括了动物对化学信号的辨别、学习和记忆。在国际上，对动物社会识别行为的研究已成为揭示动物学习和记忆的神经生物学机制的一个起点。本研究以单配制的棕色田鼠(Microtus mandarinus)为对象，探讨了它的社会识别行为，包括棕色田鼠对同种个体气味源探究格局的分析：哺乳期棕色田鼠对配偶的识别记忆；哺乳期棕色田鼠对非亲缘幼仔的识别；棕色田鼠幼仔对双亲气味的识别；棕色田鼠与沼泽田鼠(Microtus fortis)个体识别中nNOS和FOS的表达；外周注射一氧化氮合酶(NOS)抑制剂对棕色田鼠嗅觉探究及脑区FOS表达的影响。
1 棕色田鼠对同种个体气味源探究格局的分析：通过观察非动情期的棕色田鼠对麻醉的同种个体身体气味源的探究行为，包括雄鼠对麻醉雄鼠和雌鼠的嗅闻、雌鼠对麻醉雄鼠和雌鼠的嗅闻，分析了棕色田鼠在社会探究中对不同气味源的反应格局。结果表明，棕色田鼠对同种个体肛殖区和面部的嗅闻频次及嗅闻持续时间最多(P＜0.05)；对背部、腹部和肋部嗅闻的频次和持续时间较少。在对同一部位的嗅闻中，在嗅闻频次上，雄鼠对雄鼠的嗅闻、雌鼠对雌鼠的嗅闻以及雌鼠对雄鼠的嗅闻都显著多于雄鼠对雌鼠的嗅闻(P＜0.05)；在嗅闻时间上，性别组合对同一部位的探究行为没有显著性差异，但气味源与性别组合有交互作用(P＜0.05)，影响探究行为。这些结果说明棕色田鼠的肛殖区和面部是重要的气味源，而且对同一气味源的探究存在性别差异。但在非动情期，异性间的探究并不总是高于同性间的探究。
2 哺乳期棕色田鼠对配偶的识别记忆：观察了分娩后哺乳的雌性棕色田鼠与配偶分离7日、14日、21日后对配偶与陌生雄鼠的行为反应，结果发现，无论在频次和持续时间上，与配偶分离7日、14日、21日的雌鼠对配偶的攻击行为显著低于陌生雄鼠(P＜0.05)；对配偶的嗅闻及友善行为显著高于陌生雄鼠(P＜0.05)。处于哺乳前、中、后期不同阶段的雌鼠对陌生雄鼠攻击水平没有显著性差异(P＞0.05)。这些结果说明在整个哺乳期，雌鼠能够识别配偶，对配偶的记忆不因分离时间的增加而减弱。而且对陌生鼠的攻击水平不会因哺乳阶段的变化而变化，维持高水平的攻击可能与其较强的母性行为有关。
3 哺乳期棕色田鼠对非亲缘幼仔的识别：观察了产后2日(L2)、6日(L6)、10日(L10)、14日(L14)和20日(L20)的棕色田鼠母鼠对亲仔和2日、6日、10日和
14日龄非亲缘幼仔的行为反应;以及14日龄非亲缘幼仔对母鼠衔回行为的反应。
以期探讨棕色田鼠母性识别与母鼠产后时间、非亲缘幼仔日龄的关系。结果发现，
不同日龄的非亲缘幼仔在母鼠哺乳初期(LZ，L6)被攻击的频次有显著性差异
(P<0.01);不同日龄的非亲缘幼仔在母鼠哺乳中(LIO，L14)、后期(L 20)被攻
击的频次没有显著性差异。在整个哺乳期，母鼠对14日龄非亲缘幼仔的杀婴行为
最强(P<0.01)。产后不同时间的母鼠对同一日龄非亲缘幼仔的杀婴行为没有显著
差异。14日龄非亲缘幼仔与亲缘幼仔相比，对母鼠的衔回行为表现出较强的防御
性攻击(P<0 .01)。这些结果说明，棕色田鼠母鼠对幼仔存在着母性识别，而且这
种识别可能在分娩后2日就己建立，对非亲缘幼仔的杀婴行为与母性识别、幼仔
的日龄或个体大小有关，与母鼠的产后时间无关。同时，幼仔可能在14日龄已能
够识别非亲缘母鼠。
4棕色田鼠幼仔对双亲气味的识别:利用气味选择实验，观察了10日龄、14
日龄、18日龄、22日龄的棕色田鼠幼仔对亲本底物和陌生哺乳鼠底物的行为反应，
探讨了幼体对双亲气味和陌生鼠气味的识别能力。10日龄的幼仔对双亲底物与陌
生底物的偏向没有显著性差异。14日龄和18日龄的幼仔对双亲底物的访问、嗅闻
和挖掘行为均明显多于陌生底物(P<0 .05)。22日龄的幼仔对双亲底物的嗅闻和挖
掘行为明显多于陌生底物(P<0.05)。说明产后14日龄的棕色田鼠幼仔己能够辨
别双亲气味和陌生鼠气味。产后14日龄的幼仔己能睁眼，幼仔发育过程中的“睁
眼”不仅是其生理发育的一个重要阶段，而且可能在幼仔识别双亲的过程中发挥
重要作用。
5棕色田鼠与沼泽田鼠个体识别中riNOS和FOS的表达:通过居住一入侵
(resident-intruder)的行为实验和SABC免疫组织化学相结合的方法，探讨了棕色
田鼠和沼泽田鼠对入侵的陌生同种个体的识别情况。棕色田鼠和沼泽田鼠的居住
鼠与入侵鼠经过Zh的相互作用后，居住鼠对入侵鼠的探究行为没有显著性差异，
但攻击行为明显减少(P<0.05)。在定量分析前梨状皮层(PCa)、杏仁内侧核(ME)、
杏仁皮质前核(ACO)和下丘脑室旁核(PVN)的FOS和nNOS的表达后发现，
居住鼠与入侵鼠2h的相互作用后，棕色田鼠和沼泽田鼠的FOS和nNOS在前梨状
皮层、杏仁内侧核和杏仁皮质前核明显增多(P<0.01)。但棕色田鼠室旁核的nNOS
明显减少(P<0 .01)，而沼泽田鼠没有显著性差异，FOS阳性颗粒在棕色田鼠与沼
泽田鼠的室旁核都有显著增多(P<0 .01)。这些结果表明，棕色田鼠和沼泽田鼠在
领域防卫后均能够识别入侵鼠，前梨状皮层和杏仁核可能参与了这一识别过程，
nNOS在室旁核的不同表达可能
Chemical cues profoundly influence rodents' behavior, and play a significant role to animals' survival. Social relationships in rodents are dependent on social recognition. This recognition includes rodents identify, learn and remember chemical signals. International studies for animal social recognition behavior have made it a start elucidating animal learn and memory neurobiology mechanisms. This study use monogamous mandarin voles (Microtus mandarinus) as subject to investigate following social recognition behavior: Analysis of exploratory behavior pattern of mandarin voles for conspecific individual odor sources; Lactating mandarin voles recognition memory for mate; Lactating mandarin vole dams recognition for alien pups; Mandarin vole pups recognition for parental odors; Expression of nNOS and FOS in individual recognition of mandarin voles and reed voles (Microtus fortis); Effects of peripheral administration nitric oxide synthesis (NOS) inhibitor to mandarin voles olfactory exploration and F
OS expression in brain.
1. Analysis of exploratory behavior pattern of mandarin voles for conspecific individual odor sources: It was observed that exploratory behavior of anoestrus mandarin voles for different odor sources of anesthetized conspecific individuals, including males sniffling anesthetized males and females as well females sniffling anesthetized males and females. Results showed that mandarin voles displayed a greater number of investigations for anogenital region (A-R) and face than for other sources (P<0.05); Sniffling frequency and duration was lower in the regions of back, venter and flank. Sniffling frequency for the same region of males to males, females to females, females to males was higher than that of males to females (PO.05); Sniffling duration for the same region of gender arranging in parrs had no significant difference, but odor sources and gender arranging in pairs had interaction which influenced investigating behavior (P<0.05). These results indicate that A-R and face are important regions for the vo
les producing individual odor information. Although gender difference for the voles investigating the same odor source was found, yet not all investigations of heterosexual individuals are more than those of homosexual individuals in the anoestrus period.
2. Lactating mandarin voles recognition memory for mate: In a resident-intruder test, lactating mandarin voles were observed for their behavior response to strange males and their mates that were separated from females for 7days, 14days and 21 days
after females parturition. Manifested by frequency and duration, females showed a lower aggressive behavior to mates than to strange males (P<0.05), in contrast, females showed more amicability behavior and sniffling behavior to mates than to strange males (P<0.05). Maternal aggression level had no significantly difference during early, middle and late lactating periods. These results suggest that mandarin vole females are capable of forming selective mate recognition and stranger-directed aggression in the whole lactating period. Although females are in different lactating phase, yet their aggression level to strangers has no obvious change, which may be related with their strong maternal behavior.
3. Lactating mandarin vole dams recognition for alien pups: Relation on maternal recognition, postnatal time and alien pups age with infanticide in lactating mandarin voles were investigated. Mandarin vole dams in home cage during postnatal 2days (L2), 6days (L6), lOdays (L10), 14days (L14) and 20days (L20) were observed and compared for behavioral response to their own pups and 2-day-old, 6-day-old, 10-day-old and 14-day-old alien pups, While 14-day-old alien pups were observed when they were retrieved by dams. During early lactating period (L2, L4), dams infanticide to different age alien pups had significantly difference (P<0.01), and this phenomenon was not found during mid (LI4, L10) and late (L20) lactating period. Dams displayed more infanticide to 14-day-old alien old pups in the whole lac
1 棕色田鼠对同种个体气味源探究格局的分析：通过观察非动情期的棕色田鼠对麻醉的同种个体身体气味源的探究行为，包括雄鼠对麻醉雄鼠和雌鼠的嗅闻、雌鼠对麻醉雄鼠和雌鼠的嗅闻，分析了棕色田鼠在社会探究中对不同气味源的反应格局。结果表明，棕色田鼠对同种个体肛殖区和面部的嗅闻频次及嗅闻持续时间最多(P＜0.05)；对背部、腹部和肋部嗅闻的频次和持续时间较少。在对同一部位的嗅闻中，在嗅闻频次上，雄鼠对雄鼠的嗅闻、雌鼠对雌鼠的嗅闻以及雌鼠对雄鼠的嗅闻都显著多于雄鼠对雌鼠的嗅闻(P＜0.05)；在嗅闻时间上，性别组合对同一部位的探究行为没有显著性差异，但气味源与性别组合有交互作用(P＜0.05)，影响探究行为。这些结果说明棕色田鼠的肛殖区和面部是重要的气味源，而且对同一气味源的探究存在性别差异。但在非动情期，异性间的探究并不总是高于同性间的探究。
2 哺乳期棕色田鼠对配偶的识别记忆：观察了分娩后哺乳的雌性棕色田鼠与配偶分离7日、14日、21日后对配偶与陌生雄鼠的行为反应，结果发现，无论在频次和持续时间上，与配偶分离7日、14日、21日的雌鼠对配偶的攻击行为显著低于陌生雄鼠(P＜0.05)；对配偶的嗅闻及友善行为显著高于陌生雄鼠(P＜0.05)。处于哺乳前、中、后期不同阶段的雌鼠对陌生雄鼠攻击水平没有显著性差异(P＞0.05)。这些结果说明在整个哺乳期，雌鼠能够识别配偶，对配偶的记忆不因分离时间的增加而减弱。而且对陌生鼠的攻击水平不会因哺乳阶段的变化而变化，维持高水平的攻击可能与其较强的母性行为有关。
3 哺乳期棕色田鼠对非亲缘幼仔的识别：观察了产后2日(L2)、6日(L6)、10日(L10)、14日(L14)和20日(L20)的棕色田鼠母鼠对亲仔和2日、6日、10日和
14日龄非亲缘幼仔的行为反应;以及14日龄非亲缘幼仔对母鼠衔回行为的反应。
以期探讨棕色田鼠母性识别与母鼠产后时间、非亲缘幼仔日龄的关系。结果发现，
不同日龄的非亲缘幼仔在母鼠哺乳初期(LZ，L6)被攻击的频次有显著性差异
(P<0.01);不同日龄的非亲缘幼仔在母鼠哺乳中(LIO，L14)、后期(L 20)被攻
击的频次没有显著性差异。在整个哺乳期，母鼠对14日龄非亲缘幼仔的杀婴行为
最强(P<0.01)。产后不同时间的母鼠对同一日龄非亲缘幼仔的杀婴行为没有显著
差异。14日龄非亲缘幼仔与亲缘幼仔相比，对母鼠的衔回行为表现出较强的防御
性攻击(P<0 .01)。这些结果说明，棕色田鼠母鼠对幼仔存在着母性识别，而且这
种识别可能在分娩后2日就己建立，对非亲缘幼仔的杀婴行为与母性识别、幼仔
的日龄或个体大小有关，与母鼠的产后时间无关。同时，幼仔可能在14日龄已能
够识别非亲缘母鼠。
4棕色田鼠幼仔对双亲气味的识别:利用气味选择实验，观察了10日龄、14
日龄、18日龄、22日龄的棕色田鼠幼仔对亲本底物和陌生哺乳鼠底物的行为反应，
探讨了幼体对双亲气味和陌生鼠气味的识别能力。10日龄的幼仔对双亲底物与陌
生底物的偏向没有显著性差异。14日龄和18日龄的幼仔对双亲底物的访问、嗅闻
和挖掘行为均明显多于陌生底物(P<0 .05)。22日龄的幼仔对双亲底物的嗅闻和挖
掘行为明显多于陌生底物(P<0.05)。说明产后14日龄的棕色田鼠幼仔己能够辨
别双亲气味和陌生鼠气味。产后14日龄的幼仔己能睁眼，幼仔发育过程中的“睁
眼”不仅是其生理发育的一个重要阶段，而且可能在幼仔识别双亲的过程中发挥
重要作用。
5棕色田鼠与沼泽田鼠个体识别中riNOS和FOS的表达:通过居住一入侵
(resident-intruder)的行为实验和SABC免疫组织化学相结合的方法，探讨了棕色
田鼠和沼泽田鼠对入侵的陌生同种个体的识别情况。棕色田鼠和沼泽田鼠的居住
鼠与入侵鼠经过Zh的相互作用后，居住鼠对入侵鼠的探究行为没有显著性差异，
但攻击行为明显减少(P<0.05)。在定量分析前梨状皮层(PCa)、杏仁内侧核(ME)、
杏仁皮质前核(ACO)和下丘脑室旁核(PVN)的FOS和nNOS的表达后发现，
居住鼠与入侵鼠2h的相互作用后，棕色田鼠和沼泽田鼠的FOS和nNOS在前梨状
皮层、杏仁内侧核和杏仁皮质前核明显增多(P<0.01)。但棕色田鼠室旁核的nNOS
明显减少(P<0 .01)，而沼泽田鼠没有显著性差异，FOS阳性颗粒在棕色田鼠与沼
泽田鼠的室旁核都有显著增多(P<0 .01)。这些结果表明，棕色田鼠和沼泽田鼠在
领域防卫后均能够识别入侵鼠，前梨状皮层和杏仁核可能参与了这一识别过程，
nNOS在室旁核的不同表达可能
Chemical cues profoundly influence rodents' behavior, and play a significant role to animals' survival. Social relationships in rodents are dependent on social recognition. This recognition includes rodents identify, learn and remember chemical signals. International studies for animal social recognition behavior have made it a start elucidating animal learn and memory neurobiology mechanisms. This study use monogamous mandarin voles (Microtus mandarinus) as subject to investigate following social recognition behavior: Analysis of exploratory behavior pattern of mandarin voles for conspecific individual odor sources; Lactating mandarin voles recognition memory for mate; Lactating mandarin vole dams recognition for alien pups; Mandarin vole pups recognition for parental odors; Expression of nNOS and FOS in individual recognition of mandarin voles and reed voles (Microtus fortis); Effects of peripheral administration nitric oxide synthesis (NOS) inhibitor to mandarin voles olfactory exploration and F
OS expression in brain.
1. Analysis of exploratory behavior pattern of mandarin voles for conspecific individual odor sources: It was observed that exploratory behavior of anoestrus mandarin voles for different odor sources of anesthetized conspecific individuals, including males sniffling anesthetized males and females as well females sniffling anesthetized males and females. Results showed that mandarin voles displayed a greater number of investigations for anogenital region (A-R) and face than for other sources (P<0.05); Sniffling frequency and duration was lower in the regions of back, venter and flank. Sniffling frequency for the same region of males to males, females to females, females to males was higher than that of males to females (PO.05); Sniffling duration for the same region of gender arranging in parrs had no significant difference, but odor sources and gender arranging in pairs had interaction which influenced investigating behavior (P<0.05). These results indicate that A-R and face are important regions for the vo
les producing individual odor information. Although gender difference for the voles investigating the same odor source was found, yet not all investigations of heterosexual individuals are more than those of homosexual individuals in the anoestrus period.
2. Lactating mandarin voles recognition memory for mate: In a resident-intruder test, lactating mandarin voles were observed for their behavior response to strange males and their mates that were separated from females for 7days, 14days and 21 days
after females parturition. Manifested by frequency and duration, females showed a lower aggressive behavior to mates than to strange males (P<0.05), in contrast, females showed more amicability behavior and sniffling behavior to mates than to strange males (P<0.05). Maternal aggression level had no significantly difference during early, middle and late lactating periods. These results suggest that mandarin vole females are capable of forming selective mate recognition and stranger-directed aggression in the whole lactating period. Although females are in different lactating phase, yet their aggression level to strangers has no obvious change, which may be related with their strong maternal behavior.
3. Lactating mandarin vole dams recognition for alien pups: Relation on maternal recognition, postnatal time and alien pups age with infanticide in lactating mandarin voles were investigated. Mandarin vole dams in home cage during postnatal 2days (L2), 6days (L6), lOdays (L10), 14days (L14) and 20days (L20) were observed and compared for behavioral response to their own pups and 2-day-old, 6-day-old, 10-day-old and 14-day-old alien pups, While 14-day-old alien pups were observed when they were retrieved by dams. During early lactating period (L2, L4), dams infanticide to different age alien pups had significantly difference (P<0.01), and this phenomenon was not found during mid (LI4, L10) and late (L20) lactating period. Dams displayed more infanticide to 14-day-old alien old pups in the whole lac
引文
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