进化发展心理学：发展中的人类本性

进化发展心理学研究社会能力与认知能力发展背后的基因与环境机制,以及基因与环境机制在特定社会文化中的表现。进化发展心理学探讨的主要问题与设计到的主要概念包括：（1）自然选择在毕生发展的早期阶段发挥作用;（2）进化发展心理学特别重视适应在婴儿期及儿童期发展中的作用;（3）儿童期的延长有助于儿童学习人类社会的复杂性;（4）发展背景的方法可以用来理解成人外显行为的进化起源;（5）基因与环境因素相互影响约束个体发展;（6）认知能力与行为活动敏感于早期环境,但二者发展也具有可塑性     

A mathematical model of the evolution of individual differences in developmental plasticity arising through parental bet‐hedging

Children vary in the extent to which their development is shaped by particular experiences (e.g. maltreatment, social support). This variation raises a question: Is there no single level of plasticity that maximizes biological fitness? One influential hypothesis states that when different levels of plasticity are optimal in different environmental states and the environment fluctuates unpredictably, natural selection may favor parents producing offspring with varying levels of plasticity. The current article presents a mathematical model assessing the logic of this hypothesis – specifically, it examines what conditions are required for natural selection to favor parents to bet‐hedge by varying their offspring's plasticity. Consistent with existing theory from biology, results show that between‐individual variation in plasticity cannot evolve when the environment only varies across space. If, however, the environment varies across time, selection can favor differential plasticity, provided fitness effects are large (i.e. variation in individuals’ plasticity is correlated with substantial variation in fitness). Our model also generates a novel restriction: Differential plasticity only evolves when the cost of being mismatched to the environment exceeds the benefits of being well matched. Based on mechanistic considerations, we argue that bet‐hedging by varying offspring plasticity, if it were to evolve, would be more likely instantiated via epigenetic mechanisms (e.g. pre‐ or postnatal developmental programming) than genetic ones (e.g. mating with genetically diverse partners). Our model suggests novel avenues for testing the bet‐hedging hypothesis of differential plasticity, including empirical predictions and relevant measures. We also discuss several ways in which future work might extend our model.     

Biological Sensitivity to Context

ABSTRACT— Conventional views suggest that exaggerated biological reactivity to stress is a harmful vestige of an evolutionary past in which threats to survival were more prevalent and severe. Recent evidence, however, indicates that effects of high reactivity on behavior and health are bivalent rather than univalent in character, exerting both risk-augmenting and risk-protective effects depending on the context. These observations suggest that heightened stress reactivity may reflect increased biological sensitivity to context, with potential for negative health effects under conditions of adversity and for positive effects under conditions of support. From an evolutionary perspective, the developmental plasticity of the stress-response systems, along with their structured, context-dependent effects, suggests that variation in these systems has been adaptively patterned to increase the capacity of children to match their stress-response profiles to anticipated developmental environments. Taken together, these theoretical perspectives generate a novel hypothesis: that there is a curvilinear, U-shaped relation between early exposures to adversity and the development of stress-reactive profiles, with high-reactivity phenotypes disproportionately emerging within both highly stressful and highly protected early social environments.     

Functions and mental representation: the theoretical role of representations and its real nature

Representations are not only used in our folk-psychological explanations of behaviour, but are also fruitfully postulated, for example, in cognitive science. The mainstream view in cognitive science maintains that our mind is a representational system. This popular view requires an understanding of the nature of the entities they are postulating. Teleosemantic theories face this challenge, unpacking the normativity in the relation of representation by appealing to the teleological function of the representing state. It has been argued that, if intentionality is to be explained in teleological terms, then the function of a state cannot depend on its phylogenetical history, given the metaphysical possibility of a duplicate of an intentional being that lacks an evolutionary history (Swampman). In this paper, I present a method to produce, according to our current knowledge in genetic engineering, human-like individuals who are not the product of natural selection in the required sense. This variation will be used to shed light on the main replies that have been offered in the literature to the Swampman thought experiment. I argue that these replies are not satisfactory: representations should better not depend on natural selection. I conclude that a non-etiological notion of function is to be preferred for characterizing the relation of representation.     

On the epigenetic evolution of species-specific perception: the developmental manifold concept

As evolution proceeds, natural selection operates to preserve adaptive phenotypes. The adaptive phenotypes are produced during the course of individual development. Thus, natural selection favors animals that have had a particular developmental history, including, but not restricted to, their genes. This point is demonstrated by the importance of normally occurring embryonic sensory stimulation in the development of instinctive perception in two species of ducklings.     

Précis of Evolution in Four Dimensions

In his theory of evolution, Darwin recognized that the conditions of life play a role in the generation of hereditary variations, as well as in their selection. However, as evolutionary theory was developed further, heredity became identified with genetics, and variation was seen in terms of combinations of randomly generated gene mutations. We argue that this view is now changing, because it is clear that a notion of hereditary variation that is based solely on randomly varying genes that are unaffected by developmental conditions is an inadequate basis for evolutionary theories. Such a view not only fails to provide satisfying explanations of many evolutionary phenomena, it also makes assumptions that are not consistent with the data that are emerging from disciplines ranging from molecular biology to cultural studies. These data show that the genome is far more responsive to the environment than previously thought, and that not all transmissible variation is underlain by genetic differences. In Evolution in Four Dimensions (2005) we identify four types of inheritance (genetic, epigenetic, behavioral, and symbol-based), each of which can provide variations on which natural selection will act. Some of these variations arise in response to developmental conditions, so there are Lamarckian aspects to evolution. We argue that a better insight into evolutionary processes will result from recognizing that transmitted variations that are not based on DNA differences have played a role. This is particularly true for understanding the evolution of human behavior, where all four dimensions of heredity have been important.     

Behavioral genetic variation, adaptation and maladaptation: an evolutionary perspective

Evolutionary psychology seeks to understand the adaptive, evolved nature of humans by considering the forces of natural selection that gave rise to it. Individual humans also exhibit maladaptation, however, and some of the variation in maladaptive conditions is heritable. An evolutionary perspective can shed light on these phenomena too. Recent work suggests that developmental imprecision importantly affects the fitness of organisms and some of the genetic influences have been identified. Both theory and evidence that developmental imprecision underlies human cognitive maladaptation has begun to accumulate, although the manner in which the developing brain is affected remains largely unknown.     

Erroneous choice and foregone gains in hoarding gray jays (Perisoreus canadensis)

Under the conventional assumption that natural selection favors choice behavior that maximizes some fitness-related currency, a forager making repeated binary choices should consistently choose the more valuable option. Under the alternative assumption that natural selection favors choice behavior that minimizes costly errors, erroneous choice is not only expected but is expected to be common when the cost of errors is low. This cost depends on the potential rate of return: the higher this rate, the smaller the cost of choosing the less valuable option. When this rate is very high, a forager may err frequently and yet forego no appreciable fitness gain. Errors should thus be more common when interruptions to foraging are shorter. Our experimental results supported this prediction: gray jays chose the less valuable option more frequently when subjected to shorter interruptions (experimentally imposed delays to access to food rewards). This tendency is consistent with the idea that an adaptive decision-making process may routinely produce errors, not because errors are in some way adaptive but because their fitness cost is minimal, particularly when delays are short. From a proximate perspective, this tendency to commit errors more frequently following shorter delays may be due to constraints on the jays’ information-processing capacity. In general, choice behavior should be viewed as the joint byproduct of adaptive decision making and cognitive constraints. Received: 8 June 2000 / Accepted after revision: 21 October 2000     

The origins of extraversion: joint effects of facultative calibration and genetic polymorphism

The origins of variation in extraversion are largely mysterious. Recent theories and some findings suggest that personality variation can be orchestrated by specific genetic polymorphisms. Few studies, however, have examined an alternative hypothesis that personality traits are facultatively calibrated to variations in other phenotypic features, and none have considered how these distinct processes may interact in personality determination. Since physical strength and physical attractiveness likely predicted the reproductive payoffs of extraverted behavioral strategies over most of human history, it was theorized that extraversion is calibrated to variation in these characteristics. Confirming these predicted patterns, strength and attractiveness together explained a surprisingly large fraction of variance in extraversion across two studies--effects that were independent of variance explained by an androgen receptor gene polymorphism. These novel findings initially support an integrative model wherein facultative calibration and specific genetic polymorphisms operate in concert to determine personality variation.     

A cGMP-dependent protein kinase gene, foraging, modifies habituation-like response decrement of the giant fiber escape circuit in Drosophila

The Drosophila giant fiber jump-and-flight escape response is a model for genetic analysis of both the physiology and the plasticity of a sensorimotor behavioral pathway. We previously established the electrically induced giant fiber response in intact tethered flies as a model for habituation, a form of nonassociative learning. Here, we show that the rate of stimulus-dependent response decrement of this neural pathway in a habituation protocol is correlated with PKG (cGMP-Dependent Protein Kinase) activity and foraging behavior. We assayed response decrement for natural and mutant rover and sitter alleles of the foraging (for) gene that encodes a Drosophila PKG. Rover larvae and adults, which have higher PKG activities, travel significantly farther while foraging than sitters with lower PKG activities. Response decrement was most rapid in genotypes previously shown to have low PKG activities and sitter-like foraging behavior. We also found differences in spontaneous recovery (the reversal of response decrement during a rest from stimulation) and a dishabituation-like phenomenon (the reversal of response decrement evoked by a novel stimulus). This electrophysiological study in an intact animal preparation provides one of the first direct demonstrations that PKG can affect plasticity in a simple learning paradigm. It increases our understanding of the complex interplay of factors that can modulate the sensitivity of the giant fiber escape response, and it defines a new adult-stage phenotype of the foraging locus. Finally, these results show that behaviorally relevant neural plasticity in an identified circuit can be influenced by a single-locus genetic polymorphism existing in a natural population of Drosophila.     

Interval timing as an emergent learning property

Interval timing in operant conditioning is the learned covariation of a temporal dependent measure such as wait time with a temporal independent variable such as fixed-interval duration. The dominant theories of interval timing all incorporate an explicit internal clock, or "pacemaker," despite its lack of independent evidence. The authors propose an alternative, pacemaker-free view that demonstrates that temporal discrimination can be explained by using only 2 assumptions: (a) variation and selection of responses through competition between reinforced behavior and all other, elicited, behaviors and (b) modulation of the strength of response competition by the memory for recent reinforcement. The model departs radically from existing timing models: It shows that temporal learning can emerge from a simple dynamic process that lacks a periodic time reference such as a pacemaker.     

When is it Selectively Advantageous to Have True Beliefs? Sandwiching the Better Safe than Sorry Argument

Several philosophers have argued that natural selection will favor reliable belief formation; others have been more skeptical. These traditional approaches to the evolution of rationality have been either too sketchy or else have assumed that phenotypic plasticity can be equated with having a mind. Here I develop a new model to explore the functional utility of belief and desire formation mechanisms, and defend the claim that natural selection favors reliable inference methods in a broad, but not universal, range of circumstances.     

Word cues affect detection but not localization responses

Many theories assume that preknowledge of an upcoming target helps visual selection. In those theories, a top-down set can alter the salience of the target, such that attention can be deployed to the target more efficiently and responses are faster. Evidence for this account stems from visual search studies in which the identity of the upcoming target is cued in advance. In five experiments, we show that top-down knowledge affects the speed with which a singleton target can be detected but not the speed with which it can be localized. Furthermore, we show that these results are independent of the mode of responding (manual or saccadic) and are not due to a ceiling effect. Our results suggest that in singleton search, top-down information does not affect visual selection but most likely does affect response selection. We argue that such an effect is found only when information from different dimensions needs to be integrated to generate a response and that this is the case in singleton detection tasks but not in other singleton search tasks.     

Evolution and ontogeny of stress response to social challenges in the human child

The stress response systems of the human child are highly sensitive to social challenges. Because stress hormones can have negative developmental and health consequences, this presents an evolutionary paradox: Why would natural selection have favored mechanisms that elevate stress hormone levels in response to psychosocial stimuli? Two complementary hypotheses are considered: (a) maladaptation to the novelty of chronic stress in social environments, and (b) adaptive neural reorganization that facilitates the ontogeny of social competencies. Data on salivary cortisol, morbidity, and social environment from an 18 year study of child health in a rural community on the island of Dominica are examined from the perspective of these alternative hypotheses. Results indicate that difficult family environments and traumatic events are associated with elevated cortisol levels and higher morbidity. The long-term effects of traumatic early experiences on cortisol profiles are complex and indicate domain-specific effects, with normal recovery from physical stressors, but heightened response to negative-affect social challenges.     

Animal cognition + optimal choice = behavior: A review of adaptive behavior and learning, 2nd Ed., by J. E. R. Staddon

Staddon discusses a vast array of topics in comparative psychology in this book. His view is that adaptive behavior in most cases is the result of optimal choice acting on an animal's knowledge about the world. Staddon refers to this as a functional teleonomic approach inasmuch as it attempts to understand an animal's behavior in terms of goals. He builds mathematical models based on this idea that are designed to reproduce specific sets of empirical observations, usually qualitatively. A natural consequence of Staddon's approach is that many models are developed, each of which applies to a specific set of observations. An alternative to functional teleonomy is a functional approach that builds on prior principles. In most cases, this approach favors a single‐theory account of behavior. Prior principles can be understood as functional stand‐ins for antecedent material causes, which means that these accounts are closer to mechanistic theories than are goal‐based teleonomic accounts. An ontological perspective, referred to as supervenient realism, is a means of understanding the relationship between functional theories and the material world. According to this perspective, the algorithmic operation of a successful functional theory may be understood to supervene on the material operation of the nervous system.     

Zinc-induced peripheral anosmia and behavioral responses to novelty in mice: a quantitative-genetic analysis

Adult male mice were made anosmic by intranasal flushing with a 5% zinc sulfate solution. Twelve behavioral variables were measured in treated as well as saline-irrigated control animals placed in a novel environment. The genetic underpinnings and the genotype-treatment interactions with regard to these behaviors were analyzed in a classical Mendelian cross between the inbred strains C57BL/6 and DBA/2 and in a full 4 X 4 diallel cross, replicated five times, between these strains and strains C3H/St an CPB-K. Based on the hypothesis of an evolutionary history of directional selection for a well-balanced information-processing system, one might expect directional dominance for decrease in exploration after anosmization. Although decreases were found for several behavioral phenotypes, only few and relatively unimportant genotype-treatment interactions were present. This absence of any kind of genetic variation for behavioral change after anosmization points to an extremely strong directional selection which has eliminated all less favorable alleles. The findings support the hypothesis of directional selection for an efficient olfactory information-processing system.     

Genetic architecture of olfactory discriminative avoidance conditioning in Drosophila melanogaster

Recent claims to have demonstrated associative learning ability in Drosophila melanogaster raise questions about the adaptive significance of behavioral modifiability of this species. In a strain survey and a 9 X 9 half diallel cross study of olfactory discriminative avoidance conditioning, a low narrow heritability and strong directional dominance or heterosis controlling nonrandom phenotypic variation were found. Furthermore, the predicted inbreeding depression and asymmetrical response to bidirectional genetic selection were both observed. The genetic architecture revealed in these experiments is consistent with a close association between this conditioning phenotype and evolutionary fitness. Predictions from this interpretation to the nature of new mutations have been confirmed, and a possible role for conditioning in courtship behavior has been identified.     

Application of a Multidimensional Nested Logit Model to Multiple-Choice Test Items

Nested logit models have been presented as an alternative to multinomial logistic models for multiple-choice test items (Suh and Bolt in Psychometrika 75:454–473, 2010) and possess a mathematical structure that naturally lends itself to evaluating the incremental information provided by attending to distractor selection in scoring. One potential concern in attending to distractors is the possibility that distractor selection reflects a different trait/ability than that underlying the correct response. This paper illustrates a multidimensional extension of a nested logit item response model that can be used to evaluate such distinctions and also defines a new framework for incorporating collateral information from distractor selection when differences exist. The approach is demonstrated in application to questions faced by a university testing center over whether to incorporate distractor selection into the scoring of its multiple-choice tests. Several empirical examples are presented.     

Probability matching in choice under uncertainty: Intuition versus deliberation

Gaissmaier and Schooler (2008) [Gaissmaier, W., & Schooler, L. J. (2008). The smart potential behind probability matching. Cognition, 109, 416-422] argue that probability matching, which has traditionally been viewed as a decision making error, may instead reflect an adaptive response to environments in which outcomes potentially follow predictable patterns. In choices involving monetary stakes, we find that probability matching persists even when it is not possible to identify or exploit outcome patterns and that many “probability matchers” rate an alternative strategy (maximizing) as superior when it is described to them. Probability matching appears to reflect a mistaken intuition that can be, but often is not, overridden by deliberate consideration of alternative choice strategies.     

On the strategic modulation of the time course of facilitation and inhibition of return

In studies of exogenous attentional orienting, response times for targets at previously cued locations are often longer than those for targets at previously uncued locations. This effect is known widely as inhibition of return (IOR). There has been debate as to whether IOR can be observed in discrimination as well as detection tasks. The experiments reported here confirm that IOR can be observed when target discrimination is required and that the cue-target interval at which IOR is observed is often longer in discrimination than in detection tasks. The results also demonstrate that the later emergence of IOR is related to perceptual discrimination rather than to response selection differences between discrimination and detection tasks. More difficult discrimination tasks lengthen the SOA at which IOR emerges. In contrast, increasing task difficulty by adding a distractor to the location opposite the target shortens the SOA at which IOR emerges. Together, the results reveal an adaptive interaction between exogenous and endogenous attentional systems, in which the action of the orienting (exogenous) system is modulated endogenously in accord with task demands.