利他行为的遗传基础: 来自定量遗传学和分子遗传学的证据

doi:10.3724/SP.J.1042.2022.01574

摘要/Abstract

摘要：

利他行为是指个体耗损自身资源帮助他人的行为,对群体和个人而言具有重要意义。在现实生活中,利他行为具有极大的个体差异,而遗传在其中起着关键作用。近年来,基于定量遗传学和分子遗传学两类研究方法,研究者关注利他行为的遗传基础。首先,采用定量遗传学研究方法探讨利他行为的遗传率,已有的大量研究证实利他行为确实受遗传所影响,同时遗传率估计在不同研究中有所差异（0~0.87）,利他的遗传程度可能受年龄、测量方法、环境等多种因素影响。其次,基于分子遗传学研究方法,研究者发现了四类利他候选基因,包括多巴胺受体基因、5-羟色胺转运体基因、催产素受体基因、和加压素受体基因,以上发现证实一些基因位点与利他行为具有一定的相关性。同时不容忽视的是,无论是在定量遗传学研究,还是分子遗传学研究中,环境已被证实不仅会影响基因遗传率,同时也在基因对行为的影响过程中发挥着关键作用。一方面基因与环境相关联,二者共同影响利他行为,即基因-环境相关,共有三种基因-环境相关类型,包括被动型、唤起型、和主动型;另一方面基因的效应受环境影响,即差别易感性模型,外部环境以“更好或更坏”的方式影响着易感基因携带者的利他行为发展,这在催产素受体基因和多巴胺受体基因中发现了大量的基因与环境对利他的交互作用。目前研究存在利他候选基因选择具有推测性,现有结论不一致等问题,未来研究需拓展神经生物系统探索,注重全基因组研究、元分析和机制探索,和进行系统的环境干预实践。

关键词: 利他行为, 基因, 基因-环境相关, 差别易感性

Abstract:

Altruistic behavior is a behavior that benefits others at a cost to oneself, which is of great significance to groups and individuals. People differ significantly in their everyday altruistic behaviors, which is partly influenced by genetic factors. Recently, researchers have focused on the role of genes in altruistic behavior.
First, the heritability of altruistic behavior was explored based on the method of quantitative genetics. A large number of studies have confirmed that altruistic behavior is indeed affected by heredity, and heritability estimates vary among studies (0~0.87). The heritability of altruistic behavior may be influenced by factors such as age, the method of measurement, and some environmental factors (e.g., culture, family environments). Second, based on molecular genetic research, researchers have found four categories of altruism-related candidate genes, including dopamine receptor genes, serotonin transporter genes, oxytocin receptor genes, and vasopressin receptor genes. These findings confirmed that altruistic behavior was correlated with some gene loci. Taken together, both quantitative and molecular genetic studies have provided abundant genetic evidence of altruistic behavior.
Furthermore, according to the aforementioned studies, the environment has been proven not only to affect the heritability of altruistic behavior but also to play a key role in the influence of genes on altruistic behavior in both quantitative and molecular genetic studies. On the one hand, genotype is associated with an environment that jointly influences altruistic behavior, known as gene-environment correlation. There are three types of gene-environment correlations: passive, evocative and active. On the other hand, the effect of genetics on altruistic behavior is influenced by the environment, known as the differential susceptibility model; that is, the environment affects the development of the altruistic behavior of susceptibility gene carriers in a manner of “strengthening or weakening.” Accordingly, a large number of findings regarding the interactions between genes and the environment on altruism have been found in oxytocin receptor genes and dopamine receptor genes.
The current studies have some problems, such as speculative selection of altruistic candidate genes and inconsistent conclusions. Future research needs to expand on and further explore the effect of neurobiological systems on altruistic behavior, which may focus on genome-wide research, meta-analysis, mechanism exploration, and systematic environmental intervention practice.

Key words: altruistic behavior, gene, gene-environment correlation, differential susceptibility

中图分类号:

李海虹, 尚思源, 谢晓非. (2022). 利他行为的遗传基础: 来自定量遗传学和分子遗传学的证据. 心理科学进展 , 30(7), 1574-1588.

LI Haihong, SHANG Siyuan, XIE Xiaofei. (2022). The role of genes in altruistic behavior: Evidence from quantitative genetics and molecular genetics. Advances in Psychological Science, 30(7), 1574-1588.

图/表 2

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候选基因	分类	位点/基因型	环境因素	利他测量工具或范式	结果
多巴胺受体基因	G	DRD₄(4R, 7R)	--	改编的NEO人格问卷(Revised NEO Personality Inventory, NEO- PI-R)	4R基因携带者比7R基因携带者更利他(Anacker et al., 2013)
	G	DRD₄(4R, 7R)	--	无私量表	4R基因携带者比7R基因携带者更利他(Bachner-Melman et al., 2005)
	G	DRD₄(4R, 7R)	--	最后通牒博弈	4R基因携带者比7R基因携带者更利他(Reuter et al., 2013)
	G×E	DRD₄(2R, 7R)	宗教信仰	志愿活动意愿	启动宗教信仰, 易感性基因(2R、7R)携带者志愿活动意愿更高, 未携带者不受宗教信仰启动影响(Sasaki et al., 2013)
	G×E	DRD₄(2R, 7R)	攻击性家庭环境	PTM中文版(Prosocial Tendencies Measure, PTM)的利他分维度	在高攻击性家庭环境中, 易感性基因(2R、7R)携带者利他倾向更低; 在低攻击性家庭环境中, 非易感性基因携带者利他倾向更低(王云强等, 2020)
	G×E	DRD₄(4R)	宗教信仰, 性别	独裁者博弈	4R基因携带者, 宗教与利他无显著关联; 非4R基因携带者, 基督教徒男性比非基督教徒更利他(Jiang et al., 2015)
	G×E	DRD₄(7R)	父母教养方式	SDQ问卷	父母教养方式和DRD₄-7R基因对儿童自评利他行为交互作用不显著(Bersted, 2016)
5-羟色胺转运体基因	G	5-HTTLPR (LL, SS, LS)	--	人际反应指针量表(Davis’ Interpersonal Reactivity Index, IRI)	5-HTTLPR基因与特质共情无显著关联(Huetter et al., 2020)
5-羟色胺转运体基因	G	5-HTTLPR (LL, SS, LS)	--	帮助量表	L基因携带者比S基因携带者更帮助他人(Stoltenberg et al., 2013)
催产素受体基因	G	rs53576 (GG, AA, AG)	--	顾客导向量表(customer orientation scale)	SNP (rs53576)GG型个体比A等位基因携带者(即AG, AA)更愿意帮助客户(Verbeke et al., 2013)
	G	rs1042778, rs2268490, rs237887	--	独裁者博弈社会价值取向问卷	3种SNP基因与两个利他任务有显著关联, 其中rs1042778相关性最高(Israel et al., 2009)
	G	rs53576 (GG, AA, AG)	--	观察者观察一对伴侣求助互动无声视频	SNP (rs53576)GG型个体比A等位基因携带者(即AG, AA)更利他, 且可被观察者识别(Kogan et al., 2011)
	G	rs7632287, rs237887, rs2268493, rs237897, rs1042778, rs53576, rs2254298, rs75775, rs4686302,	--	独裁者博弈信任游戏	9种SNP与两个利他任务无显著关联(Apicella et al., 2010)
	G×E	rs53576 (GG, AA, AG)	感知的环境威胁	慈善行为	SNP (rs53576)A等位基因(AA、AG)携带者, 环境威胁感知越高慈善行为越少; GG型个体, 环境威胁感知与慈善行为无显著关联(Poulin et al., 2012)
	G×E	rs2254298 (GG, AA, AG)	性别	PTM中文版	男性个体SNP (rs2254298)G等位基因(AG、GG)携带者更利他, 女性个体三种基因携带者利他倾向无显著差异(Shang et al., 2017)
	G×E	rs13316193 (CC, TT, CT)	接受者身份、帮助成本	帮助情境	当接受者是外群体且帮助成本较高时, SNP (rs13316193)C等位基因(CT/CC)携带者比TT型个体帮助倾向更高; 低成本以及接受者是内群体时, 不同基因型面对高/低帮助成本帮助倾向无差异 (Wu & Su, 2018)
候选基因	分类	位点/基因型	环境因素	利他测量工具或范式	结果
加压素受体基因	G	AVPR1aRS3 (327~343bp, 308~325bp)	--	独裁者博弈	长版基因携带者比短版基因携带者更利他(Knafo et al., 2008)
	G	AVPR1aRS3 (327bp)	--	独裁者博弈	携带目标等位基因的儿童更不利他(Avinun et al., 2011)
	G	AVPR1b的rs28373064 (GG, AA, AG)	--	改编的亲社会倾向量表(revised Prosocial Tendencies Measure, PTM-R), 包含利他分量表	AVPR1b的SNP (rs28373064)G等位基因携带者比A等位基因携带者更利他(Wu et al., 2015)

候选基因	分类	位点/基因型	环境因素	利他测量工具或范式	结果
多巴胺受体基因	G	DRD₄(4R, 7R)	--	改编的NEO人格问卷(Revised NEO Personality Inventory, NEO- PI-R)	4R基因携带者比7R基因携带者更利他(Anacker et al., 2013)
	G	DRD₄(4R, 7R)	--	无私量表	4R基因携带者比7R基因携带者更利他(Bachner-Melman et al., 2005)
	G	DRD₄(4R, 7R)	--	最后通牒博弈	4R基因携带者比7R基因携带者更利他(Reuter et al., 2013)
	G×E	DRD₄(2R, 7R)	宗教信仰	志愿活动意愿	启动宗教信仰, 易感性基因(2R、7R)携带者志愿活动意愿更高, 未携带者不受宗教信仰启动影响(Sasaki et al., 2013)
	G×E	DRD₄(2R, 7R)	攻击性家庭环境	PTM中文版(Prosocial Tendencies Measure, PTM)的利他分维度	在高攻击性家庭环境中, 易感性基因(2R、7R)携带者利他倾向更低; 在低攻击性家庭环境中, 非易感性基因携带者利他倾向更低(王云强等, 2020)
	G×E	DRD₄(4R)	宗教信仰, 性别	独裁者博弈	4R基因携带者, 宗教与利他无显著关联; 非4R基因携带者, 基督教徒男性比非基督教徒更利他(Jiang et al., 2015)
	G×E	DRD₄(7R)	父母教养方式	SDQ问卷	父母教养方式和DRD₄-7R基因对儿童自评利他行为交互作用不显著(Bersted, 2016)
5-羟色胺转运体基因	G	5-HTTLPR (LL, SS, LS)	--	人际反应指针量表(Davis’ Interpersonal Reactivity Index, IRI)	5-HTTLPR基因与特质共情无显著关联(Huetter et al., 2020)
5-羟色胺转运体基因	G	5-HTTLPR (LL, SS, LS)	--	帮助量表	L基因携带者比S基因携带者更帮助他人(Stoltenberg et al., 2013)
催产素受体基因	G	rs53576 (GG, AA, AG)	--	顾客导向量表(customer orientation scale)	SNP (rs53576)GG型个体比A等位基因携带者(即AG, AA)更愿意帮助客户(Verbeke et al., 2013)
	G	rs1042778, rs2268490, rs237887	--	独裁者博弈社会价值取向问卷	3种SNP基因与两个利他任务有显著关联, 其中rs1042778相关性最高(Israel et al., 2009)
	G	rs53576 (GG, AA, AG)	--	观察者观察一对伴侣求助互动无声视频	SNP (rs53576)GG型个体比A等位基因携带者(即AG, AA)更利他, 且可被观察者识别(Kogan et al., 2011)
	G	rs7632287, rs237887, rs2268493, rs237897, rs1042778, rs53576, rs2254298, rs75775, rs4686302,	--	独裁者博弈信任游戏	9种SNP与两个利他任务无显著关联(Apicella et al., 2010)
	G×E	rs53576 (GG, AA, AG)	感知的环境威胁	慈善行为	SNP (rs53576)A等位基因(AA、AG)携带者, 环境威胁感知越高慈善行为越少; GG型个体, 环境威胁感知与慈善行为无显著关联(Poulin et al., 2012)
	G×E	rs2254298 (GG, AA, AG)	性别	PTM中文版	男性个体SNP (rs2254298)G等位基因(AG、GG)携带者更利他, 女性个体三种基因携带者利他倾向无显著差异(Shang et al., 2017)
	G×E	rs13316193 (CC, TT, CT)	接受者身份、帮助成本	帮助情境	当接受者是外群体且帮助成本较高时, SNP (rs13316193)C等位基因(CT/CC)携带者比TT型个体帮助倾向更高; 低成本以及接受者是内群体时, 不同基因型面对高/低帮助成本帮助倾向无差异 (Wu & Su, 2018)
候选基因	分类	位点/基因型	环境因素	利他测量工具或范式	结果
加压素受体基因	G	AVPR1aRS3 (327~343bp, 308~325bp)	--	独裁者博弈	长版基因携带者比短版基因携带者更利他(Knafo et al., 2008)
	G	AVPR1aRS3 (327bp)	--	独裁者博弈	携带目标等位基因的儿童更不利他(Avinun et al., 2011)
	G	AVPR1b的rs28373064 (GG, AA, AG)	--	改编的亲社会倾向量表(revised Prosocial Tendencies Measure, PTM-R), 包含利他分量表	AVPR1b的SNP (rs28373064)G等位基因携带者比A等位基因携带者更利他(Wu et al., 2015)