读心的比较研究：非人灵长类与人类在心理理论上的异同点及其解释

doi:10.3724/SP.J.1042.2022.02540

摘要/Abstract

摘要：

利用非人灵长类开展比较研究是探索心理理论本质、起源和进化动因的重要途经。大量实证研究显示非人灵长类可以表现出一些基础心理理论成分, 包括联合注意、意图理解和一级观点采择, 这表明心理理论在灵长类动物的进化谱系中具有进化连续性。然而, 非人灵长类无法像人类一样达成二级观点采择和错误信念理解, 潜心智化假说、替代表征假说、最小限度心理理论假说和意识关系假说试图对此差异进行解释。尽管视角与具体内容不同, 但这些假说都强调, 非人灵长类无法对其他个体关于外部世界的心理信念形成表征, 信念表征是非人灵长类与人类读心能力的重要分界点。未来研究需要重点澄清共享意图、语言机制以及社会生态环境因素与心理理论产生的相互影响作用。

关键词: 非人灵长类, 心理理论, 联合注意, 观点采择, 错误信念

Abstract:

Theory of mind refers to the ability of individuals to speculate about other people's mental states and make causal explanations for their behaviors. Comparative studies using non-human primates is an important way to explore the essence, origin and evolution of theory of mind. This article reviews a large number of empirical studies and demonstrate that nonhuman primates exhibit some basic theory of mind components, including joint attention, goal understanding and level-1 perspective-taking. Nonhuman primates can follow or attract the attention of other individuals, understand the behavior of other individuals based on potential goals and intentions, and judge whether other individuals perceive something from their perspective.

The paper explains the above conclusions from the perspective of evolutionary continuity hypothesis. Species that appeared earlier in the primate evolutionary lineage have been shown to have basic theory of mind. Phylogenetic reconstruction proved that theory of mind has deep evolutionary roots in primates. Research from neuroscience is consistent with this view. Brain imaging studies have shown that non-human primates and humans rely on common brain regions for mind-reading tasks. On the one hand, there are obvious anatomical and connectivity similarities between these parts of the human and non-human primate brains. On the other hand, these areas of the human brain have undergone significant expansion over the course of evolution. So theory of mind has evolutionary continuity across primate lineages. The similarity between nonhuman primates and humans in theory of mind stem from shared descent.

The paper also concludes that non-human primates are unable to achieve second-order perspective selection and false belief understanding as humans do. The submentalizing hypothesis, vicarious representation hypothesis, minimal theory of mind hypothesis and awareness relations hypothesis attempt to explain the differences. Although specific views and perspectives vary, these theories all emphasize that there is a primary representational system before a mature theory of mind is mastered by the individual, and these representational systems lay the foundation for the development of a complete theory of mind. However, non-human primates only have a primary representation system, and they are unable to form representations of other individuals' psychological beliefs about the external world. Belief representation is an important dividing point between non-human primates and humans in theory of mind. Unfortunately, there is no single hypothesis that can explain all the current empirical research. One possibility is that higher-level theory of mind depend on many different sub-processing mechanisms. Different task paradigms depend on different processing mechanisms, thus resulting in seemingly contradictory performance of non-human primates in different experimental modes. The differences and related explanations between non-human primates and human psychological theories in theory of mind need to be tested more fully in future studies.

For humans, theory of mind is a psychological mechanism that plays an important role in both competitive and cooperative scenarios. Non-human primates, on the other hand, seem to be more likely to initiate mind-reading in competitive situations, and competitive activities lead to more pronounced use of theory of mind. They are more likely to perceive competitive intentions from others than cooperative intentions or shared intentions. One reasonable speculation is that during human evolution, specific social pressures led humans to develop sensitivity to shared intentions. Theory of mind is further improved when individuals are able to establish common goals and coordinate with each other for common goals. Future research should focus on this hypothesis, and further explore the influence of shared intention on the development of primate theory of mind through empirical research.

The relationship between language and theory of mind is another topic that needs to be explored in the future. On the one hand, a higher level of theory of mind allows the individual to step into the perspective of the person he is communicating with, allowing him to understand the true meaning of the sender of the information senders. So the improvement of mind-reading ability in humans may have been a key shift in the evolution of language. Theory of mind is an important basis for the realization of complex language. On the other hand, language system provides a necessary tool for mental representation system. When represented in language, subjects can encode complex nested recursive relationships. Future research should focus on the relationship between the characteristics of non-human primates' theory of mind and their communication systems, and seek enlightenment for revealing the mysteries of human theory of mind and language evolution through comparative studies of different non-human primates.

Furthermore, as an important social cognitive feature, theory of mind is bound to be closely related to the social ecological environment of a species. In the future, more comparative studies on differences can enable researchers to explore the relationship between theory of mind and social attributes, and also enable researchers to test whether each sub-component of theory of mind has different social driving factors, providing important basis for understanding the evolutionary mechanism of theory of mind.

Key words: non-human primates, theory of mind, joint attention, perspective-taking, false belief

中图分类号:

B844

殷融. (2022). 读心的比较研究：非人灵长类与人类在心理理论上的异同点及其解释. 心理科学进展 , 30(11), 2540-2557.

YIN Rong. (2022). Comparative studies of mind reading: Similarities and differences in theory of mind between non-human primates and humans and corresponding explanations. Advances in Psychological Science, 30(11), 2540-2557.

图/表 4

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研究者	实验内容	实验结果	指向结论
Castellano-Navarro 等(2021)	检验了日本猕猴(Japanese Macaques)在不同情境中对人类目光的反应：威胁场景中的实验员是一个猕猴不熟悉的人, 当他逐渐走近猕猴时猕猴会把他视为威胁; 合作情境中的实验员是一个经常为猕猴提供食物的人, 实验员用手拿着食物, 头或身体扭向不同方向。	在威胁场景下, 相比实验员凝视其他方向, 如果实验员直接凝视猕猴, 猕猴会表现出更强烈的逃离倾向; 合作场景下, 猕猴会根据实验员的注视方向来调整自身位置, 让自己进入实验员视野。	猕猴对人类目光很敏感, 它们可以根据情境灵活地领会人类注视的意义, 并做出不同行为反应。
Bettl和Rosati (201920192019); MacLean和 Hare (2012)	恒河猴(Rhesus)/黑猩猩目睹实验员从背面凝视一个箱子, 在无遮挡情况下, 箱子是透明的; 在有遮挡情况下, 被试看不到箱子背面, 研究者纪录被试的行为反应。	相比无遮挡情况, 在遮挡情况下被试有更大比率会移动到实验员所在方位, 观察箱子内部。	恒河猴/黑猩猩会积极追踪其他个体的目光, 努力识别其他个体到底看到了什么。
Sánchez-Amaro 等(2020)	以白颊长臂猿(Hoolock Gibbons)和银白长臂猿(Silvery Gibbons)为被试, 允许长臂猿在两份食物之间选择, 两份食物与实验员具有不同方位关系。	长臂猿会更倾向于选择实验员背对方位的食物或与实验员头部朝向不一致方位的食物。	长臂猿可以利用头部或身体朝向等线索来理解其他个体的注意力状态。
Aychet等(2020); Deshpande等(2018)	在实验中允许红头白眉猴(Red- capped Mangabey)/印度野生冠毛猕猴(Bonnet Macaques)在实验员四周自由移动, 实验员手拿食物奖励, 以不同的身体角度面对动物。	动物被试会主动移动身体, 让自己进入实验员视野; 只有在同实验员相互注视时, 被试才表现出乞食动作; 当食物与实验员目光方向不一致时, 被试会优先移动到实验员面前与对方保持视觉接触。	非人灵长类会评估其他个体的注意状态, 并在有需要时主动吸引对方注意。

研究者	实验内容	实验结果	指向结论
Castellano-Navarro 等(2021)	检验了日本猕猴(Japanese Macaques)在不同情境中对人类目光的反应：威胁场景中的实验员是一个猕猴不熟悉的人, 当他逐渐走近猕猴时猕猴会把他视为威胁; 合作情境中的实验员是一个经常为猕猴提供食物的人, 实验员用手拿着食物, 头或身体扭向不同方向。	在威胁场景下, 相比实验员凝视其他方向, 如果实验员直接凝视猕猴, 猕猴会表现出更强烈的逃离倾向; 合作场景下, 猕猴会根据实验员的注视方向来调整自身位置, 让自己进入实验员视野。	猕猴对人类目光很敏感, 它们可以根据情境灵活地领会人类注视的意义, 并做出不同行为反应。
Bettl和Rosati (201920192019); MacLean和 Hare (2012)	恒河猴(Rhesus)/黑猩猩目睹实验员从背面凝视一个箱子, 在无遮挡情况下, 箱子是透明的; 在有遮挡情况下, 被试看不到箱子背面, 研究者纪录被试的行为反应。	相比无遮挡情况, 在遮挡情况下被试有更大比率会移动到实验员所在方位, 观察箱子内部。	恒河猴/黑猩猩会积极追踪其他个体的目光, 努力识别其他个体到底看到了什么。
Sánchez-Amaro 等(2020)	以白颊长臂猿(Hoolock Gibbons)和银白长臂猿(Silvery Gibbons)为被试, 允许长臂猿在两份食物之间选择, 两份食物与实验员具有不同方位关系。	长臂猿会更倾向于选择实验员背对方位的食物或与实验员头部朝向不一致方位的食物。	长臂猿可以利用头部或身体朝向等线索来理解其他个体的注意力状态。
Aychet等(2020); Deshpande等(2018)	在实验中允许红头白眉猴(Red- capped Mangabey)/印度野生冠毛猕猴(Bonnet Macaques)在实验员四周自由移动, 实验员手拿食物奖励, 以不同的身体角度面对动物。	动物被试会主动移动身体, 让自己进入实验员视野; 只有在同实验员相互注视时, 被试才表现出乞食动作; 当食物与实验员目光方向不一致时, 被试会优先移动到实验员面前与对方保持视觉接触。	非人灵长类会评估其他个体的注意状态, 并在有需要时主动吸引对方注意。

研究者	实验内容	实验结果	指向结论
Buttelmann等(2007)	黑猩猩观察实验员用不寻常的身体部位(比如头或脚)控制一个开关, 随后将黑猩猩置于同样的情境中, 观察黑猩猩行为。	如果实验员双手没有被占用, 随后黑猩猩也会更容易模仿实验员的动作; 但如果实验员双手被其他东西所占用, 那么随后黑猩猩则会更倾向于用手去操纵开关, 它们会“理性模仿”而不是完全复制。	非人灵长类被试能理解其他个体的真正目标, 可以透过他们的“表面动作”认知到其行为意图。
Rochat等(2008)	猕猴先看到实验员从障碍物后伸手去拿特定目标物; 障碍移除后, 实验员使用不同姿势去拿目标物。	当障碍移除后, 如果实验员仍然使用相同姿势去拿目标物, 猕猴会对他注视更长的时间, 它们对实验员这一“没必要”的动作表示不解。	非人灵长类被试能理解其他个体的真正目标, 可以透过他们的“表面动作”认知到其行为意图。
Buttelmann等(2012)	猿类被试看到实验员尝试打开一个上了锁的箱子(测试箱)：在一种情境下, 实验员已先后打开其他几个箱子并把箱子里的食物给予被试; 而在另一种情境下, 实验员也已先后尝试打开其他几个箱子, 但没成功, 不过每次动作结束后他会把口袋里的食物给予被试。	箱子获取食物情境下, 黑猩猩和倭黑猩猩被试会在实验员摆弄测试箱时表现得更加耐心, 它们可以理解在这种情境中实验员的意图是打开箱子将食物喂给它们, 而另一种情境中实验员只是想尝试打开箱子。	非人灵长类可以综合经验、动作与环境线索, 对其他个体的行为意图进行推测。
Canteloup和Meunier (2017)	以汤基猕猴(Tonkean Macaque)为被试, 实验员手拿葡萄干, 他反复做出要给猕猴葡萄干的动作, 但猕猴实际得不到食物。在一种情况下, 猕猴得不到葡萄干的原因是实验员在戏弄它, 在另一种情况下, 猕猴得不到葡萄干的原因是葡萄干塞不进笼子的进食口。	在故意戏弄情境中, 猕猴会对实验员和他手里的食物注视更长时间, 它们做出的乞食动作会更少, 但会更频繁地尝试抓取实验员手中的食物。猕猴能够区分一个人是不愿意给它们食物还是无法给它们食物。	非人灵长类可以综合经验、动作与环境线索, 对其他个体的行为意图进行推测。
Yamamoto等(2012)	两只黑猩猩被透明或不透明的屏障隔开, 其中一只黑猩猩的旁边有一杯果汁, 但它需要合适的工具才能够取到果汁, 这只猩猩隔着屏障伸出手向另一只黑猩猩求助, 后者身边有许多工具, 它要选择将哪一个工具递给求助者。	当被试可以看到求助者那一侧的情况时, 它会倾向于将合适的工具(弯柄木棍)递给求助者; 但当被试无法看到求助者那一侧的情况时, 它将各种工具递给求助者的可能性相当。	非人灵长类可以综合环境和动作信息理解其他个体是否有得到某个物品的意愿, 而不仅仅依据单一的动作或空间关系做出判断。
Drayton和Santos (2014)	以棕色卷尾猴(Cebus Apella)为被试, 实验员做出各种动作, 试图在众多物品中, 获得一个够不到的目标物, 卷尾猴可以拿到目标物。	卷尾猴会把实验员想要的目标物给他, 而不是其他物品; 甚至由于障碍阻隔, 实验员只能将手伸向目标物的方向时, 大部分卷尾猴也能做到这一点; 然而如果实验员转过身去, 从背后伸手对着目标物, 卷尾猴就不会表现出传递倾向。	非人灵长类可以综合环境和动作信息理解其他个体是否有得到某个物品的意愿, 而不仅仅依据单一的动作或空间关系做出判断。

研究者	实验内容	实验结果	指向结论
Buttelmann等(2007)	黑猩猩观察实验员用不寻常的身体部位(比如头或脚)控制一个开关, 随后将黑猩猩置于同样的情境中, 观察黑猩猩行为。	如果实验员双手没有被占用, 随后黑猩猩也会更容易模仿实验员的动作; 但如果实验员双手被其他东西所占用, 那么随后黑猩猩则会更倾向于用手去操纵开关, 它们会“理性模仿”而不是完全复制。	非人灵长类被试能理解其他个体的真正目标, 可以透过他们的“表面动作”认知到其行为意图。
Rochat等(2008)	猕猴先看到实验员从障碍物后伸手去拿特定目标物; 障碍移除后, 实验员使用不同姿势去拿目标物。	当障碍移除后, 如果实验员仍然使用相同姿势去拿目标物, 猕猴会对他注视更长的时间, 它们对实验员这一“没必要”的动作表示不解。	非人灵长类被试能理解其他个体的真正目标, 可以透过他们的“表面动作”认知到其行为意图。
Buttelmann等(2012)	猿类被试看到实验员尝试打开一个上了锁的箱子(测试箱)：在一种情境下, 实验员已先后打开其他几个箱子并把箱子里的食物给予被试; 而在另一种情境下, 实验员也已先后尝试打开其他几个箱子, 但没成功, 不过每次动作结束后他会把口袋里的食物给予被试。	箱子获取食物情境下, 黑猩猩和倭黑猩猩被试会在实验员摆弄测试箱时表现得更加耐心, 它们可以理解在这种情境中实验员的意图是打开箱子将食物喂给它们, 而另一种情境中实验员只是想尝试打开箱子。	非人灵长类可以综合经验、动作与环境线索, 对其他个体的行为意图进行推测。
Canteloup和Meunier (2017)	以汤基猕猴(Tonkean Macaque)为被试, 实验员手拿葡萄干, 他反复做出要给猕猴葡萄干的动作, 但猕猴实际得不到食物。在一种情况下, 猕猴得不到葡萄干的原因是实验员在戏弄它, 在另一种情况下, 猕猴得不到葡萄干的原因是葡萄干塞不进笼子的进食口。	在故意戏弄情境中, 猕猴会对实验员和他手里的食物注视更长时间, 它们做出的乞食动作会更少, 但会更频繁地尝试抓取实验员手中的食物。猕猴能够区分一个人是不愿意给它们食物还是无法给它们食物。	非人灵长类可以综合经验、动作与环境线索, 对其他个体的行为意图进行推测。
Yamamoto等(2012)	两只黑猩猩被透明或不透明的屏障隔开, 其中一只黑猩猩的旁边有一杯果汁, 但它需要合适的工具才能够取到果汁, 这只猩猩隔着屏障伸出手向另一只黑猩猩求助, 后者身边有许多工具, 它要选择将哪一个工具递给求助者。	当被试可以看到求助者那一侧的情况时, 它会倾向于将合适的工具(弯柄木棍)递给求助者; 但当被试无法看到求助者那一侧的情况时, 它将各种工具递给求助者的可能性相当。	非人灵长类可以综合环境和动作信息理解其他个体是否有得到某个物品的意愿, 而不仅仅依据单一的动作或空间关系做出判断。
Drayton和Santos (2014)	以棕色卷尾猴(Cebus Apella)为被试, 实验员做出各种动作, 试图在众多物品中, 获得一个够不到的目标物, 卷尾猴可以拿到目标物。	卷尾猴会把实验员想要的目标物给他, 而不是其他物品; 甚至由于障碍阻隔, 实验员只能将手伸向目标物的方向时, 大部分卷尾猴也能做到这一点; 然而如果实验员转过身去, 从背后伸手对着目标物, 卷尾猴就不会表现出传递倾向。	非人灵长类可以综合环境和动作信息理解其他个体是否有得到某个物品的意愿, 而不仅仅依据单一的动作或空间关系做出判断。

研究者	实验内容	实验结果	指向结论
Drayton和Santos (2017) MacLean和Hare (2012)	在实验中要求实验员注视某个方向后表现表现出吃惊的样子, 在此之前, 实验员曾查看或没查看同一方向近处的目标物, 黑猩猩/倭黑猩猩/恒河猴在旁边观看了全过程。	在前一种情况下动物被试会用更长的时间观察实验员注视方向的远处, 被试知道实验员已获知关于近处目标物的信息, 因此它们会在远处寻找让实验员吃惊的刺激。	非人灵长类不仅仅是被动追随他者的注意状态, 它们可以从其他个体的视角出发, 理解它们能看到什么以及看不到什么。
Bräuer等(2007)	一只居于从属地位的黑猩猩和一只居于支配地位的黑猩猩相互面对面坐着, 在它们之间有两个平行放置的屏障, 一道屏障是透明的, 一道是不透明的, 实验员在每一道屏障靠近从属者那一侧都放了食物, 这样支配者就只能看到透明屏障后的食物但看不到不透明屏障后的食物, 之后允许被试进行食物选择。	当允许从属者先进行选择时, 它们倾向于优先选择不透明屏障后的食物。
MacLean等(2013)	以狐猴(Lemur)为被试, 研究者将食物放在两个人类助手面前, 一个助手看着食物, 另一个闭着眼睛头扭向其他方位, 允许狐猴选择食物。	狐猴会优先去拿不被助手注意的食物。
Bohn等(2016)	两只黑猩猩对藏在三个杯子里的食物依次做出选择, 它们之间具有信息不对称性, 被试看到两种食物被分别放在两个杯子下面, 而它的竞争对手只看到一种食物放在一个杯子下面。实验开始后黑猩猩轮流选择, 但它们无法看到对方的选择结果。	如果被试后做出选择, 它会倾向于选择自己看到但竞争对手没看到的放食物的杯子, 也就是说被试能推断出, 竞争对手选择时已经选择了它自己所看到的那个放食物的杯子。	非人灵长类可以通过观察来推断其它个体是否知道某些信息, 并据此来预测对方的行为。
Drayton和Santos (2018)	在预期违背任务实验中, 地中海猕猴看到实验员目睹一颗柠檬滚入到两个盒子中的一个, 之后实验员去打开放着柠檬的盒子或另一个空盒子。	当实验员接近空盒子时被试会注视更长时间。被试知道实验员掌握了柠檬藏在哪个盒子里, 同时还认为他应该会按照自己已掌握的信息去行动。	非人灵长类可以通过观察来推断其它个体是否知道某些信息, 并据此来预测对方的行为。
Karg等(2016)	让两只黑猩猩从不同角度观察两个尺寸完全一样的面包棍, 面包都贴在一堵障碍的同一面, 一个露出的高一些, 一个露出的矮一些。从被试的角度看, 它能看到两个面包是一样长的, 但从竞争者的角度看, 其中一个面包会更长一些。被试与竞争者轮流选择面包, 且不知道彼此的选择结果。	黑猩猩被试并没有表现出特定选择倾向, 被试无法代入竞争者的视角, 它没有推测出竞争者会先选择那个在它看来更长的面包。	黑猩猩/倭黑猩猩无法认知到, 对于同一事物自己与其他个体的观察视角可以是不同的; 也无法看出同一情境下其他个体与自己的需求是不同的, 它们不能克服自我中心倾向。
Nolte和 Call (2021)	以倭黑猩猩和黑猩猩为被试, 被试两两组合配对完成合作任务, 在一个特殊装置中, 被试要将合适的工具递给对方, 才能帮助对方(或者对方和自己一起)得到食物奖励, 双方所需要的工具是不一样的。	黑猩猩无法完成任务, 而倭黑猩猩也很难在第一时间就将合适的工具给合作伙伴, 它们倾向于认为对方与自己需要的工具是相同的。