动物使用工具行为研究进展北大核心CSCD

使用工具曾被认为是人类独有的能力,然而,在过去的50年中,学界逐渐认识到工具的使用普遍存在于整个动物界。其中,使用工具最多的类群是哺乳类、鸟类和昆虫。动物使用工具有一定目标性,然而大多数动物仅考虑当前的目标,而非长远目标。动物使用工具的行为受到环境因素和动物自身认知能力、生理特点与进化历史的影响,并可能表现出一定的个体差异。有些动物使用工具的行为是与生俱来的,然而大部分高等动物通过试错学习获得使用工具的能力。通过模仿学习,一些使用工具的行为可以传播和演化,从而在种群中广泛分布。工具的使用是动物认知领域的核心概念之一,开展动物使用工具的研究,能够加深对动物认知能力和行为进化的理解。     

Individual and social learning processes involved in the acquisition and generalization of tool use in macaques

Macaques can efficiently use several tools, but their capacity to discriminate the relevant physical features of a tool and the social factors contributing to their acquisition are still poorly explored. In a series of studies, we investigated macaques' ability to generalize the use of a stick as a tool to new objects having different physical features (study 1), or to new contexts, requiring them to adapt the previously learned motor strategy (study 2). We then assessed whether the observation of a skilled model might facilitate tool-use learning by naive observer monkeys (study 3). Results of study 1 and study 2 showed that monkeys trained to use a tool generalize this ability to tools of different shape and length, and learn to adapt their motor strategy to a new task. Study 3 demonstrated that observing a skilled model increases the observers' manipulations of a stick, thus facilitating the individual discovery of the relevant properties of this object as a tool. These findings support the view that in macaques, the motor system can be modified through tool use and that it has a limited capacity to adjust the learnt motor skills to a new context. Social factors, although important to facilitate the interaction with tools, are not crucial for tool-use learning.     

Tool use, aye-ayes, and sensorimotor intelligence

Humans, chimpanzees, capuchins and aye-ayes all display an unusually high degree of encephalization and diverse omnivorous extractive foraging. It has been suggested that the high degree of encephalization in aye-ayes may be the result of their diverse, omnivorous extractive foraging behaviors. In combination with certain forms of tool use, omnivorous extractive foraging has been hypothesized to be linked to higher levels of sensorimotor intelligence (stages 5 or 6). Although free-ranging aye-ayes have not been observed to use tools directly in the context of their extractive foraging activities, they have recently been reported to use lianas as tools in a manner that independently suggests that they may possess stage 5 or 6 sensorimotor intelligence. Although other primate species which display diverse, omnivorous extractive foraging have been tested for sensorimotor intelligence, aye-ayes have not. We report a test of captive aye-ayes' comprehension of tool use in a situation designed to simulate natural conditions. The results support the view that aye-ayes do not achieve stage 6 comprehension of tool use, but rather may use trial-and-error learning to develop tool-use behaviors. Other theories for aye-aye encephalization are considered.     

Young children spontaneously invent wild great apes’ tool-use behaviours

The variety and complexity of human-made tools are unique in the animal kingdom. Research investigating why human tool use is special has focused on the role of social learning: while non-human great apes acquire tool-use behaviours mostly by individual (re-)inventions, modern humans use imitation and teaching to accumulate innovations over time. However, little is known about tool-use behaviours that humans can invent individually, i.e. without cultural knowledge. We presented 2- to 3.5-year-old children with 12 problem-solving tasks based on tool-use behaviours shown by great apes. Spontaneous tool use was observed in 11 tasks. Additionally, tasks which occurred more frequently in wild great apes were also solved more frequently by human children. Our results demonstrate great similarity in the spontaneous tool-use abilities of human children and great apes, indicating that the physical cognition underlying tool use shows large overlaps across the great ape species. This suggests that humans are neither born with special physical cognition skills, nor that these skills have degraded due to our species’ long reliance of social learning in the tool-use domain.     

Discovering and learning tool-use for fishing honey by captive chimpanzees

Wild chimpanzees commonly use sticks to fish for termites, ants or honey. This ability seems to be socially transmitted to juveniles by their mothers across generations. In a natural environment, the limited visibility of this behavior with regards to the extent of stick's insertion and about the success or failure in fishing hinders the study of the underlying learning processes. This study explores the discovery and learning of tool use for fishing honey in an artificial hive by a group of four captive chimpanzees. The discovery of tool use was accidental and coactive. The speed with which the group of experimentally naive chimpanzees discovered and learned tool use suggests that wild chimpanzees of different populations independently discovered the fishing behavior. The alpha male and his ally learned before the subordinates. Here, trial-and-error learning was, as in monkeys, the main process allowing the acquisition of the tool-use technique. However, the observation of conspecifics allowed the orientation of the experimentation by the selection of clues. As suggested by Tomaselloet al. (1987). it is the understanding of the function of the tool,i.e. the cause-effect relations between the action of the demonstrator, the type of tool and the task to accomplish which confer to chimpanzees and advantage over monkeys.     

If at first you don't succeed… Studies of ontogeny shed light on the cognitive demands of habitual tool use

Many species use tools, but the mechanisms underpinning the behaviour differ between species and even among individuals within species, depending on the variants performed. When considering tool use ‘as adaptation’, an important first step is to understand the contribution made by fixed phenotypes as compared to flexible mechanisms, for instance learning. Social learning of tool use is sometimes inferred based on variation between populations of the same species but this approach is questionable. Specifically, alternative explanations cannot be ruled out because population differences are also driven by genetic and/or environmental factors. To better understand the mechanisms underlying routine but non-universal (i.e. habitual) tool use, we suggest focusing on the ontogeny of tool use and individual variation within populations. For example, if tool-using competence emerges late during ontogeny and improves with practice or varies with exposure to social cues, then a role for learning can be inferred. Experimental studies help identify the cognitive and developmental mechanisms used when tools are used to solve problems. The mechanisms underlying the route to tool-use acquisition have important consequences for our understanding of the accumulation in technological skill complexity over the life course of an individual, across generations and over evolutionary time.     

‘Captivity bias’ in animal tool use and its implications for the evolution of hominin technology

Animals in captive or laboratory settings may outperform wild animals of the same species in both frequency and diversity of tool use, a phenomenon here termed ‘captivity bias’. Although speculative at this stage, a logical conclusion from this concept is that animals whose tool-use behaviour is observed solely under natural conditions may be judged cognitively or physically inferior than if they had also been tested or observed under controlled captive conditions. In turn, this situation creates a potential problem for studies of the behaviour of extinct members of the human family tree—the hominins—as hominin cognitive abilities are often judged on material evidence of tool-use behaviour left in the archaeological record. In this review, potential factors contributing to captivity bias in primates (including increased contact between individuals engaged in tool use, guidance or shaping of tool-use behaviour by other tool-users and increased free time and energy) are identified and assessed for their possible effects on the behaviour of the Late Pleistocene hominin Homo floresiensis. The captivity bias concept provides one way to uncouple hominin tool use from cognition, by considering hominins as subject to the same adaptive influences as other tool-using animals.     

Tool-use for drinking water by immature chimpanzees of Mahale: prevalence of an unessential behavior

Use of leaves or sticks for drinking water has only rarely been observed during long-term study of wild chimpanzees (Pan troglodytes schweinfurthii) at Mahale. Recently, however, we observed 42 episodes of tool-use for drinking water (73 tools and two cases of using tool-sets) between 1999 and 2004. Interestingly, all of the performers were immature chimpanzees aged from 2 to 10 years. Immature chimpanzees sometimes observed the tool-using performance of others and subsequently reproduced the behavior, while adults usually paid no attention to the performance. This tool-use did not seem to occur out of necessity: (1) chimpanzees often used tools along streams where they could drink water without tools, (2) they used tools for drinking water from tree holes during the wet season when they could easily obtain water from many streams, and (3) the tool-using performance sometimes contained playful aspects. Between-site comparisons revealed that chimpanzees at drier habitats used tools for drinking water more frequently and in a more conventional manner. However, some variations could not be explained by ecological conditions. Such variations and the increase in this tool-use in recent years at Mahale strongly suggest that social learning plays an important role in the process of acquiring the behavior. We should note here that such behaviors that lack obvious benefits or necessity can be prevalent in a group.     

Toward a Choice-Based Judgment Bias Task for Horses

Judgment bias tasks for nonhuman animals are promising tools to assess emotional valence as a measure of animal welfare. In view of establishing a valid judgment bias task for horses, the present study aimed to evaluate 2 versions (go/no-go and active choice) of an auditory judgment bias task for horses in terms of acquisition learning and discrimination of ambiguous cues. Five mares and 5 stallions were randomly assigned to the 2 designs and trained for 10 trials per day to acquire different operant responses to a low-frequency tone and a high-frequency tone, respectively. Following acquisition learning, horses were tested on 4 days with 3 ambiguous-tone trials interspersed between the 10 high-tone and low-tone trials. All 5 go/no-go horses but only one active-choice horse successfully learned their task, indicating that it is more difficult to train horses on an active choice task than on a go/no-go task. During testing, however, go/no-go horses did not differentiate between the 3 different ambiguous cues, thereby making the validity of the test results questionable in terms of emotional valence.     

Observational learning of tool-use by young chimpanzees

In the current study two groups of young chimpanzees (4–6 and 8–9 years old) were given a T-bar and a food item that could only be reached by using the T-bar. Experimental subjects were given the opportunity to observe an adult using the stick as a tool to obtain the food; control subjects were exposed to the adult but were given no demonstration. Subjects in the older group did not learn to use the tool. Subjects in the younger group who were exposed to the demonstrator learned to use the stick as a tool much more readily than those who were not. None of the subjects demonstrated an ability to imitatively copy the demonstrator's precise behavioral strategies. More than simple stimulus enhancement was involved, however, since both groups manipulated the T-bar, but only experimental subjects used it in its function as a tool. Our findings complement naturalistic observations in suggesting that chimpanzee tool-use is in some sense «culturally transmitted» — though perhaps not in the same sense as social-conventional behaviors for which precise copying of conspecifics is crucial.     

Tool use by wild cebus monkeys at Santa Rosa National Park,Costa Rica

Although wild cebus monkeys have been observed to use tools, this behavior has been reported only rarely. No one has systematically examined tool use in wildCebus, and it is not known how prevalent tool use is in the species' natural repertoire. During 300 hr of observation on 21 wild capuchins (Cebus capucinus imitator) at Santa Rosa National Park in Costa Rica, 31 incidents of tool use, including eight different types of tool-use behavior, were observed. These observations indicate that tool use is a notable behavior pattern in this troop. Considering these incidents of tool use in conjunction with other reports on complex food-getting and preparation behavior byCebus suggests that tool use is a manifestation ofCebus' high behavioral adaptability. Since onlyCebus and the great apes (especially chimpanzees) have been observed to show such a diverse tool-use repertoire, to use tools so frequently, or to show such complex food-getting behavior in the wild, these observations also support the notion thatCebus and the great apes have followed a parallel evolutionary development of tool-using capacity.     

Tool use by aquatic animals

Tool-use research has focused primarily on land-based animals, with less consideration given to aquatic animals and the environmental challenges and conditions they face. Here, we review aquatic tool use and examine the contributing ecological, physiological, cognitive and social factors. Tool use among aquatic animals is rare but taxonomically diverse, occurring in fish, cephalopods, mammals, crabs, urchins and possibly gastropods. While additional research is required, the scarcity of tool use can likely be attributable to the characteristics of aquatic habitats, which are generally not conducive to tool use. Nonetheless, studying tool use by aquatic animals provides insights into the conditions that promote and inhibit tool-use behaviour across biomes. Like land-based tool users, aquatic animals tend to find tools on the substrate and use tools during foraging. However, unlike on land, tool users in water often use other animals (and their products) and water itself as a tool. Among sea otters and dolphins, the two aquatic tool users studied in greatest detail, some individuals specialize in tool use, which is vertically socially transmitted possibly because of their long dependency periods. In all, the contrasts between aquatic- and land-based tool users enlighten our understanding of the adaptive value of tool-use behaviour.     

The social learning of tool use by orangutans (Pongo pygmaeus)

Very little is known about the social learning of orangutans (Pongo pygmaeus), especially in the context of problem-solving situations such as tool use. Sixteen orangutans were presented with a rake-like tool and desirable but out-of-reach food. Eight subjects observed a human demonstrator use the tool in one way, while another eight observed the demonstrator use the tool in another way. Subjects behaved identically in the two experimental conditions, showing no effect of the type of demonstration observed. Analysis of individual learning curves suggested that a large component of individual trial-and-error learning was at work, even for two subjects who received additional trials with an orangutan demonstrator. This pattern of results suggests that subjects were paying attention to the general functional relations in the task and to the results obtained by the demonstrator, but not to the actual methods of tool use demonstrated. It is concluded that subjects in both conditions were employing emulation learning, not imitative learning.     

Data extraction from proteomics raw data: an evaluation of nine tandem MS tools using a large Orbitrap data set

In shot-gun proteomics raw tandem MS data are processed with extraction tools to produce condensed peak lists that can be uploaded to database search engines. Many extraction tools are available but to our knowledge, a systematic comparison of such tools has not yet been carried out. Using raw data containing more than 400,000 tandem MS spectra acquired using an Orbitrap Velos we compared 9 tandem MS extraction tools, freely available as well as commercial. We compared the tools with respect to number of extracted MS/MS events, fragment ion information, number of matches, precursor mass accuracies and agreement in-between tools. Processing a primary data set with 9 different tandem MS extraction tools resulted in a low overlap of identified peptides. The tools differ by assigned charge states of precursors, precursor and fragment ion masses, and we show that peptides identified very confidently using one extraction tool might not be matched when using another tool. We also found a bias towards peptides of lower charge state when extracting fragment ion data from higher resolution raw data without deconvolution. Collecting and comparing the extracted data from the same raw data allow adjusting parameters and expectations and selecting the right tool for extraction of tandem MS data.     

Tool-use and tool-making by captive, group-living orangutans (Pongo pygmaeus abelii) at an artificial termite mound

The present study examined the use and making of tools to obtain foodstuffs in artificial-mound holes by five captive, group-living Sumatran orangutans (Pongo pygmaeus abelii). Three adult orangutans frequently stripped leaves and twigs from a branch provided (tool-making), and then inserted the tool into a hole to obtain foodstuffs (tool-using). A 5-year-old female juvenile usually used the tools that adult orangutans had previously used, but rarely made tools herself. A 2-year-old male infant did not use any tools. The adult orangutans tend to leave one to several leaves at the top of the branch than to leave many leaves on the branch or to strip all leaves. It seemed likely that tools with appropriate leaves are easier to insert into holes and obtain more foodstuffs, compared with branches with many leaves or sticks without any leaves. When the orangutans were unable to insert a tool into a hole, they usually modified the tool and/or changed their tool-using technique, such as changing how they grasped the tool. These findings are discussed from the perspectives of the orangutan's behavioral flexibility regarding tool-use skills and hierarchical organization in food-processing techniques.     

Sex differences in the stone tool-use behavior of a wild population of burmese long-tailed macaques (Macaca fascicularis aurea)

We investigated sex differences in how Burmese long-tailed macaques (Macaca fascicularis aurea) used stone tools to open shelled food items along the shores of two islands in Laemson National Park, Thailand. Over a 2-week period in December 2009, we collected scan and focal samples on macaques when they were visible along the shores and mangroves. We found females used stones more often while feeding and used smaller tools than males. Females also processed sessile oysters more than males, whereas males processed unattached foods more than females. It was unclear which sex was overall more proficient at stone tool use, but males did perform significantly better at opening unattached food items with large pounding stones. Females also struck food items more times during tool-use bouts and at a faster rate, but no significant difference was found in average tool-use bout duration. Males processed foods slightly faster within a tool-use bout, but we were unable to detect a significant difference in the rate of food processing while foraging with tools. In summary females chipped open sessile oysters with an axing technique more than males, while males used larger stones to pound open unattached shelled food more often than females. Despite using pounding more than females, males also regularly utilized the axing technique on sessile oysters. Our results are the first assessment of sex differences in macaque stone tool use, providing a basis for comparison with tool use in other primates, and to nonfunctional forms of stone use in other macaques.     

Parasitic wasps learn and report diverse chemicals with unique conditionable behaviors

Parasitoids exploit numerous chemical cues to locate hosts and food. Whether they detect and learn chemicals foreign to their natural history has not been explored. We show that the parasitoid Microplitis croceipes can associate, with food or hosts, widely different chemicals outside their natural foraging encounters. When learned chemicals are subsequently detected, this parasitoid manifests distinct behaviors characteristic with expectations of food or host, commensurate with prior training. This flexibility of parasitoids to rapidly link diverse chemicals to resource needs and subsequently report them with recognizable behaviors offers new insights into their foraging adaptability, and provides a model for further dissection of olfactory learning related processes.     

Rats smell: odour-mediated local enhancement, in a vertical movement two-action test

In two experiments, hungry rats, Rattus norvegicus, were present in one side of an operant chamber while a conspecific demonstrator in the adjacent compartment moved a single lever either up or down for a food reward. During a subsequent test session, in which these rats were allowed access to the lever for the first time, all responses were rewarded regardless of their direction. In experiment 1, rats that were prevented from observing the direction of lever movement by means of a screen showed a reliable demonstrator-consistent response bias, while rats that had observed the direction of lever movement and in addition had access to any odour cues deposited on the lever did not. In experiment 2, each rat observed another rat (the ‘viewed’ demonstrator) moving a lever either up or down. They were then transferred into the test compartment of a different operant chamber in which another rat (the ‘box’ demonstrator) had moved the lever in the same direction as the viewed demonstrator or in the opposite direction. These observer rats showed a reliable preference for their box demonstrator's direction, but responded in the opposite direction to their viewed demonstrator. Taken together, the results of these experiments suggest that directional responding by rats in a vertical movement two-action test is influenced by demonstrator-deposited odour cues in addition to visual experience of a demonstrator's behaviour. Furthermore, while odour-mediated local enhancement gave rise to demonstrator-consistent responding, visual observation of a conspecific appeared to have the reverse effect. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Baboons, chimpanzees, and tools

Wild chimpanzees make and use tools with far greater frequency and variety than wild baboons. Sampling differences; differences in the sensory, motor, or cognitive capacities for skilled tool use; or environmental differences are not responsible. Fortuitous discovery of novel forms of tool behavior probably occurs as rarely among chimpanzees as among baboons. However, research on wild and captive chimpanzees reveals that the discovery is transmitted quickly among chimpanzee groups by observation learning, and thus becomes well-established in the group's behavioral repertoire. In contrast, study of captive baboons reveals that they acquire little information about tool behavior by observation. With little observation learning the behavior is not likely to be acquired by other group members since repeated independent discovery is improbable. Thus chimpanzees' more proficient tool behavior is due mainly to their greater capacity for observation learning. This suggests that the advent of hominid tool traditions was accompanied by a capacity for facile observation learning and thus, perhaps, by language ability.     

Hand preferences in captive orangutans (Pongo pygmaeus)

The strength of the evidence for population-level handedness in the great apes is a topic of considerable debate, yet there have been few studies of handedness in orangutans. We conducted a study of manual lateralization in a captive group of eight orangutans (Pongo pygmaeus) ranking the degrees of manual preference according to a defined framework. We analyzed five behavioral patterns: eat (one- and two-handed), make/modify tool, oral tool-use, and manual tool-use. Although some individuals showed significant manual preferences for one or more tasks, at the group-level both one-handed and two-handed eating, oral tool-use, and make/modify tool were ranked at level 1 (unlateralized). Manual tool-use was ranked at level 2, with four subjects demonstrating significant hand preferences, but no group-level bias to the right or left. Four subjects also showed hand specialization to the right or left across several tasks. These results are consistent with most previous studies of manual preference in orangutans. The emergence of manual lateralization in orangutans may relate to more complex manipulative tasks. We hypothesize that more challenging manual tasks elicit stronger hand preferences.