Agency Modulates the Lateral and Medial Prefrontal Cortex Responses in Belief-Based Decision Making

Many real-life decisions in complex and changing environments are guided by the decision maker’s beliefs, such as her perceived control over decision outcomes (i.e., agency), leading to phenomena like the “illusion of control”. However, the neural mechanisms underlying the “agency” effect on belief-based decisions are not well understood. Using functional imaging and a card guessing game, we revealed that the agency manipulation (i.e., either asking the subjects (SG) or the computer (CG) to guess the location of the winning card) not only affected the size of subjects’ bets, but also their “world model” regarding the outcome dependency. Functional imaging results revealed that the decision-related activation in the lateral and medial prefrontal cortex (PFC) was significantly modulated by agency and previous outcome. Specifically, these PFC regions showed stronger activation when subjects made decisions after losses than after wins under the CG condition, but the pattern was reversed under the SG condition. Furthermore, subjects with high external attribution of negative events were more affected by agency at the behavioral and neural levels. These results suggest that the prefrontal decision-making system can be modulated by abstract beliefs, and are thus vulnerable to factors such as false agency and attribution.     

It’s Not My Fault: Postdictive Modulation of Intentional Binding by Monetary Gains and Losses

Sense of agency refers to the feeling that one’s voluntary actions caused external events. Past studies have shown that compression of the subjective temporal interval between actions and external events, called intentional binding, is closely linked to the experience of agency. Current theories postulate that the experience of agency is constructed via predictive and postdictive pathways. One remaining problem is the source of human causality bias; people often make misjudgments on the causality of voluntary actions and external events depending on their rewarding or punishing outcomes. Although human causality bias implies that sense of agency can be modified by post-action information, convincing empirical findings for this issue are lacking. Here, we hypothesized that sense of agency would be modified by affective valences of action outcomes. To examine this issue, we investigated how rewarding and punishing outcomes following voluntary action modulate behavioral measures of agency using intentional binding paradigm and classical conditioning procedures. In the acquisition phase, auditory stimuli were paired with positive, neutral or negative monetary outcomes. Tone-reward associations were evaluated using reaction times and preference ratings. In the experimental session, participants performed a variant of intentional binding task, where participants made timing judgments for onsets of actions and sensory outcomes while playing simple slot games. Our results showed that temporal binding was modified by affective valences of action outcomes. Specifically, intentional binding was attenuated when negative outcome occurred, consistent with self-serving bias. Our study not only provides evidence for postdictive modification of agency, but also proposes a possible mechanism of human causality bias.     

Other faces in the mirror: a perspective on schizophrenia

A patient with schizophrenia may generate an action (whether manual or verbal), but not attribute the generation of that action to himself. We distinguish self-monitoring and attribution of agency, relating only the former to forward models and the mirror system. We suggest that alien hand experiences occur when an action progresses through hand control pathways with no record of disinhibition having been kept and is then seen but dismissed as external. Analogously, auditory pathways are active during verbal hallucinations and produce a subvocal verbal process, but since no record is kept of the words being created, they are treated as external. The subject then proceeds to confabulate, to provide an account for the agency.     

Walking on smooth or rough ground: passive control of pretarsal attachment in ants

The hymenopteran tarsus is equipped with claws and a movable adhesive pad (arolium). Even though both organs are specialised for substrates of different roughness, they are moved by the same muscle, the claw flexor. Here we show that despite this seemingly unfavourable design, the use of arolium and claws can be adjusted according to surface roughness by mechanical control. Tendon pull experiments in ants (Oecophylla smaragdina) revealed that the claw flexor elicits rotary movements around several (pre-) tarsal joints. However, maximum angular change of claws, arolium and fifth tarsomere occurred at different pulling amplitudes, with arolium extension always being the last movement. This effect indicates that arolium use is regulated non-neuronally. Arolium unfolding can be suppressed on rough surfaces, when claw tips interlock and inhibit further contraction of the claw flexor or prevent legs from sliding towards the body. To test whether this hypothesised passive control operates in walking ants, we manipulated ants by clipping claw tips. Consistent with the proposed control mechanism, claw pruning resulted in stronger arolium extension on rough but not on smooth substrates. The control of attachment by the insect claw flexor system demonstrates how mechanical systems in the body periphery can simplify centralised, neuro-muscular feedback control.     

Performance Feedback Processing Is Positively Biased As Predicted by Attribution Theory

A considerable literature on attribution theory has shown that healthy individuals exhibit a positivity bias when inferring the causes of evaluative feedback on their performance. They tend to attribute positive feedback internally (e.g., to their own abilities) but negative feedback externally (e.g., to environmental factors). However, all empirical demonstrations of this bias suffer from at least one of the three following drawbacks: First, participants directly judge explicit causes for their performance. Second, participants have to imagine events instead of experiencing them. Third, participants assess their performance only after receiving feedback and thus differences in baseline assessments cannot be excluded. It is therefore unclear whether the classically reported positivity bias generalizes to setups without these drawbacks. Here, we aimed at establishing the relevance of attributions for decision-making by showing an attribution-related positivity bias in a decision-making task. We developed a novel task, which allowed us to test how participants changed their evaluations in response to positive and negative feedback about performance. Specifically, we used videos of actors expressing different facial emotional expressions. Participants were first asked to evaluate the actors’ credibility in expressing a particular emotion. After this initial rating, participants performed an emotion recognition task and did—or did not—receive feedback on their veridical performance. Finally, participants re-rated the actors’ credibility, which provided a measure of how they changed their evaluations after feedback. Attribution theory predicts that participants change their evaluations of the actors’ credibility toward the positive after receiving positive performance feedback and toward the negative after negative performance feedback. Our results were in line with this prediction. A control condition without feedback showed that correct or incorrect performance alone could not explain the observed positivity bias. Furthermore, participants’ behavior in our task was linked to the most widely used measure of attribution style. In sum, our findings suggest that positive and negative performance feedback influences the evaluation of task-related stimuli, as predicted by attribution theory. Therefore, our study points to the relevance of attribution theory for feedback processing in decision-making and provides a novel outlook for decision-making biases.     

The evolution of armament strength: evidence for a constraint on the biting performance of claws of durophagous decapods

Performance data for the claws of six sympatric species of Cancer crabs confirmed a puzzling pattern reported previously for two other decapod crustaceans (stone crabs, Menippe mercenaria, and lobsters, Homarus americanus): Although biting forces increased, maximum muscle stresses (force per unit area) declined with increasing claw size. The negative allometry of muscle stress and the stress at a given claw size were fairly consistent within and among Cancer species despite significant differences in adult body size and relative claw size, but were not consistent among decapod genera. Therefore, claw height can be used as a reliable predictor of maximum biting force for the genus Cancer, but must be used with caution as a predictor of maximum biting force in wider evolutionary and biogeographical comparisons of decapods. The decline in maximum muscle stress with increasing claw size in Cancer crabs contrasts with the pattern in several other claw traits. Significantly, three traits that affect maximal biting force increased intraspecifically with increasing claw size: relative claw size, mechanical advantage, and sarcomere length of the closer muscle. Closer apodeme area and angle of pinnation of the closer muscle fibers varied isometrically with claw size. The concordant behavior of these traits suggests selection for higher biting forces in larger crabs. The contrast between the size dependence of muscle stress (negative allometry) and the remaining claw traits (isometry or positive allometry) strongly suggests that an as yet unidentified constraint impairs muscle performance in larger claws. The negative allometry of muscle stress in two distantly related taxa (stone crabs and lobsters) further suggests this constraint may be widespread in decapod crustaceans. The implications of this performance constraint for the evolution of claw size and the "arms-race" between decapod predators and their hard-shelled prey is discussed.     

THE DESIGN OF A BEAUTIFUL WEAPON: COMPENSATION FOR OPPOSING SEXUAL SELECTION ON A TRAIT WITH TWO FUNCTIONS

Male fiddler crabs, genus Uca, have one greatly enlarged claw with which they court females and threaten and fight other males. Longer claws are more effective signals but are thought to be less effective weapons because the relative closing force at the tip of the claw decreases with claw length. We studied claw morphology and fighting in Uca terpsichores and Uca beebei and found a mechanism that may resolve opposing selection for signaling and fighting ability. When males fought they delivered gripping forces not at the tips but at the tubercles on the inner margins of their claws’ fingers. As claws grow, these tubercles remain relatively close to the apex of the gape. Consequently, the mechanical advantage that governs the forces that can be delivered at these tubercles decreases only slightly with increasing claw length allowing the claw to be an effective signal and a powerful weapon. Animal weapons are exceptionally diverse in form and detail of armature and the causes of this diversity are poorly understood. We suggest that the designs of weapons may often reflect compensatory patterns of growth and placement of armature that enhances the weapon's overall utility for multiple uses in competition for mates.     

Testosterone, cortisol, and mood in a sports team competition

In humans, hormonal responses to winning/losing and their relationships to mood and status change have mostly been examined in individual athletic competitions. In this study, the salivary testosterone (T) and cortisol (C) and mood responses to a real match between two professional basketball teams were investigated. Data about individuals' contributions to outcome, performance appraisal, and attribution of outcome to internal/external factors were also collected. Results did not show statistically significant different T and C responses depending on the outcome. Negative mood was significantly enhanced, especially in the losers, while winners showed a better appraisal of team performance and a more internal attribution. T response did not show a significant relationship with mood changes, but it correlated positively with the "score/time playing" ratio, an indicator of individual participation in the outcome. Furthermore, T response correlated negatively with external attribution in winners and positively in losers. These results indicate that in a real, highly competitive situation, T changes are not directly a response to the outcome, but rather to the contribution the individual makes to it and to the causes he attributes.     

Neural Substrates for Judgment of Self-Agency in Ambiguous Situations

The sense of agency is the attribution of oneself as the cause of one’s own actions and their effects. Accurate agency judgments are essential for adaptive behaviors in dynamic environments, especially in conditions of uncertainty. However, it is unclear how agency judgments are made in ambiguous situations where self-agency and non-self-agency are both possible. Agency attribution is thus thought to require higher-order neurocognitive processes that integrate several possibilities. Furthermore, neural activity specific to self-attribution, as compared with non-self-attribution, may reflect higher-order critical operations that contribute to constructions of self-consciousness. Based on these assumptions, the present study focused on agency judgments under ambiguous conditions and examined the neural correlates of this operation with functional magnetic resonance imaging. Participants performed a simple but demanding agency-judgment task, which required them to report on whether they attributed their own action as the cause of a visual stimulus change. The temporal discrepancy between the participant’s action and the visual events was adaptively set to be maximally ambiguous for each individual on a trial-by-trial basis. Comparison with results for a control condition revealed that the judgment of agency was associated with activity in lateral temporo-parietal areas, medial frontal areas, the dorsolateral prefrontal area, and frontal operculum/insula regions. However, most of these areas did not differentiate between self- and non-self-attribution. Instead, self-attribution was associated with activity in posterior midline areas, including the precuneus and posterior cingulate cortex. These results suggest that deliberate self-attribution of an external event is principally associated with activity in posterior midline structures, which is imperative for self-consciousness.     

Relationships among morphology, clinging performance and habitat use in Liolaemini lizards

The central tenet of ecomorphological theory holds that different ecological requirements lead to different organismal designs (morphology). Here, we studied the relationships between performance (interlocking grasping) and forelimb morphological traits in species of lizards that exploit different structural habitats in a phylogenetic context. The performance (measured by the maximum force of clinging to substrate) was measured on different substrate types. After phylogenetically informed analyses, we found that arboreal and saxicolous species showed stronger resistance to mechanical traction in all substrates when compared to generalists and sand dweller lizards. These species showed a positive relationship between forelimb dimensions (humerus length and length of claw of toe 5) and maximum force exerted, on the contrary, hand width, claw height (CH) of digits III and IV and claw length of toe 4 showed a negative relationship. In addition, we observed a partial positive correlation between CH and maximal cling force on rough surfaces, but not on smooth surfaces.     

Size,but not sex,predicts pinch force and exoskeleton mechanical properties in crayfish of the genus Faxonius

Studies of animal weaponry and defensive structures rarely take into consideration their underlying mechanical properties. We measured the compressive strength and thickness of the exoskeleton of the claw (chela) in two North American crayfish species, Faxonius virilis and F. limosus. We performed similar measures on the carapace, a body region not directly involved in agonistic contests. Males of both species generated significantly stronger maximum pinch forces than females. However, these differences can be attributed to differences in claw size between the sexes. The thickness (ultrastructure) of the claw exoskeleton was a significant predictor of its compressive strength and likely explained the difference in compressive strength we observed between the two species. Neither claw thickness nor claw compressive strength was correlated with maximum pinch force. Additionally, we found that crayfish body size was a strong predictor of carapace compressive strength and thickness, whereas sex was not. The claw had greater compressive strength and thickness than the corresponding values for the carapace. Our study shows that the mechanical properties of the crayfish exoskeleton are largely a function of size and highlights the need to integrate mechanical properties into studies of animal morphology and performance.     

The causal pie model: an epidemiological method applied to evolutionary biology and ecology

A general concept for thinking about causality facilitates swift comprehension of results, and the vocabulary that belongs to the concept is instrumental in cross‐disciplinary communication. The causal pie model has fulfilled this role in epidemiology and could be of similar value in evolutionary biology and ecology. In the causal pie model, outcomes result from sufficient causes. Each sufficient cause is made up of a “causal pie” of “component causes”. Several different causal pies may exist for the same outcome. If and only if all component causes of a sufficient cause are present, that is, a causal pie is complete, does the outcome occur. The effect of a component cause hence depends on the presence of the other component causes that constitute some causal pie. Because all component causes are equally and fully causative for the outcome, the sum of causes for some outcome exceeds 100%. The causal pie model provides a way of thinking that maps into a number of recurrent themes in evolutionary biology and ecology: It charts when component causes have an effect and are subject to natural selection, and how component causes affect selection on other component causes; which partitions of outcomes with respect to causes are feasible and useful; and how to view the composition of a(n apparently homogeneous) population. The diversity of specific results that is directly understood from the causal pie model is a test for both the validity and the applicability of the model. The causal pie model provides a common language in which results across disciplines can be communicated and serves as a template along which future causal analyses can be made.     

Use of Stable Isotopes to Investigate Keratin Deposition in the Claw Tips of Ducks

Stable isotopes derived from the claws of birds could be used to determine the migratory origins of birds if the time periods represented in excised sections of claws were known. We investigated new keratin growth in the claws of adult female Lesser Scaup (Aythya affinis) by estimating the equilibration rates of stable isotopes (δ ¹³C, δ ¹⁵N, and δ ²H) from the breeding grounds into 1 mm claw tips. We sampled birds on their breeding ground through time and found that it took approximately 3–3.5 months for isotope values in most claw tips to equilibrate to isotope values that reflected those present in the environment on their breeding grounds. Results from this study suggest that isotopes equilibrate slowly into claw tips of Lesser Scaup, suggesting isotopes could potentially be used to determine the wintering grounds of birds. We suggest using controlled feeding experiments or longitudinal field investigations to understand claw growth and isotopic equilibration in claw tips. Such information would be valuable in ascertaining whether claw tips can be used in future studies to identify the migratory origins of birds.     

When the choice is ours: context and agency modulate the neural bases of decision-making

The option to choose between several courses of action is often associated with the feeling of being in control. Yet, in certain situations, one may prefer to decline such agency and instead leave the choice to others. In the present functional magnetic resonance imaging (fMRI) study, we provide evidence that the neural processes involved in decision-making are modulated not only by who controls our choice options (agency), but also by whether we have a say in who is in control (context). The fMRI results are noteworthy in that they reveal specific contributions of the anterior frontomedian cortex (viz. BA 10) and the rostral cingulate zone (RCZ) in decision-making processes. The RCZ is engaged when conditions clearly present us with the most choice options. BA 10 is engaged in particular when the choice is completely ours, as well as when it is completely up to others to choose for us which in turn gives rise to an attribution of control to oneself or someone else, respectively. After all, it does not only matter whether we have any options to choose from, but also who decides on that.     

Major claws make male fiddler crabs more conspicuous to visual predators: a test using human observers

One of the possible costs of the male fiddler crabs enlarged claw can be conspicuousness to predators. This hypothesis was tested using human observers as a model of visual predators. In the European fiddler crab, Uca tangeri Eydoux, the males' major claw is white contrasting with the orange-brownish colour of the carapace and of the feeding claw, and the mudflat background. The following morphotypes were created from close-up photographs taken in nature using an image processing software: male, male without claw, female, female with enlarged claw, male with enlarged claw of the same colour of the feeding claw, male with 75% sized claw, male with 50% sized claw. These morphotypes were then presented in a randomised order to students, using a psychology test software, which allows the measurement of response time in msec. The subjects were allowed to look at the images for an unlimited amount of time, until they detected the individual or until they decided to pass on to another image. Backgrounds (i.e. mudflat picture) without individuals were also presented as a control. Male crabs were detected significantly sooner than females. When we compared males with the claw removed with females with an enlarged claw added, the pattern is reversed and the latter are detected significantly faster. Thus, the enlarged claw seems to be the key feature that makes the individuals more conspicuous. Size and colour seem to be the main aspects of the claw's conspicuousness. The data of these experiments support the initial prediction of males being more conspicuous than females because of their enlarged claw. The possible costs and benefits of this trait, related to predation, are discussed.     

Autoradiographic observations on developing and growing claws of reptiles

Alibardi, L. 2010. Autoradiographic observations on developing and growing claws of reptiles. —Acta Zoologica (Stockholm) 91 : 233–241 The present qualitative autoradiographic analysis aims to present the main features of morphogenesis and growth of claws in reptiles. Lizard embryos treated with tritiated thymidine reveal that epidermal cell proliferation in terminal digits is prevalent in the dorsal side and gives origin to the curved unguis of the claw. Less proliferation occurs in the ventral side of the digit tip where the concave sub‐unguis is derived. Adult claws of a turtle show that thymidine‐labelled cells are present along most of the epidermis of the claw, especially at the claw tip. Also, injection of tritiated histidine and proline, indicating active protein synthesis, confirm autoradiographic labelling along most of the epidermis of claws, in particular at the apical tip. The present study indicates that proximal matrix regions, as have been described in mammalian nails, are absent in reptiles. This pattern of claw growth probably derives from that of terminal digital scales. In fact reptilian (and avian) claws are formed from a modification of scales, a different condition from that present in mammals.     

Dogs’ Eavesdropping from People’s Reactions in Third Party Interactions

Eavesdropping involves the acquisition of information from third-party interactions, and can serve to indirectly attribute reputation to individuals. There is evidence on eavesdropping in dogs, indicating that they can develop a preference for people based on their cooperativeness towards others. In this study, we tested dogs’ eavesdropping abilities one step further. In a first experiment, dogs could choose between cooperative demonstrators (the donors) who always gave food to an approaching third person (the beggar); here, the only difference between donors was whether they received positive or negative reactions from the beggar (through verbal and gestural means). Results showed that dogs preferentially approached the donor who had received positive reactions from the beggar. By contrast, two different conditions showed that neither the beggar’s body gestures nor the verbal component of the interaction on their own were sufficient to affect the dogs’ preferences. We also ran two further experiments to test for the possibility of dogs’ choices being driven by local enhancement. When the donors switched places before the choice, dogs chose at random. Similarly, in a nonsocial condition in which donors were replaced by platforms, subjects chose at chance levels. We conclude that dogs’ nonrandom choices in the present protocol relied on the simultaneous presence of multiple cues, such as the place where donors stood and several features of the beggar’s behavior (gestural and verbal reactions, and eating behavior). Nonetheless, we did not find conclusive evidence that dogs discriminated the donors by their physical features, which is a prerequisite of reputation attribution.     

A microscopic study of the marmoset claw and nail

W. E. Le Gros Clark concluded in his study of the problem of the primate claw that the essential difference between claw and nail is the presence of a terminal matrix associated with a deep layer in the claw, whereas neither terminal matrix nor deep layer exists in the nail. He demonstrated that the marmoset claw (which tips every digit except the hallux) has a thin deep layer and a recognizable terminal matrix. The present paper reports and discusses evidence that the marmoset nail also has a deep layer and terminal matrix. Although the importance of these structures in the claw is not disputed, it appears that these can no longer be considered absolute differences between claw and nail. On the basis of this evidence, it cannot be claimed that the presence of the deep layer and terminal matrix determines the distinctive shape of the claw as opposed to the nail.     

Case-based ethics education: the impact of cause complexity and outcome favorability on ethicality

case-based learning has been used across multiple disciplines, including ethics education, as an effective instructional tool. However, the value of case-based learning in ethics education has varied widely regarding case quality. Case content may significantly impact the ability of case-based ethics education to promote knowledge acquisition and knowledge transfer to future situations requiring ethical decision-making. This study examined two critical areas of ethical case content-causes and outcomes. Complexity of described causes and outcome favorability were manipulated in two ethical cases used during an ethics education course. Results suggest that including information in case studies reflecting clear, simple key causes and negative outcomes results in better ethical sensemaking and ethical decision-making. Implications regarding case content and case-based ethics education are explored.     

Future altruism: Social discounting of delayed rewards

Social discounting assesses an individual's willingness to forgo an outcome for the self in lieu of a larger outcome for someone else. The purpose of the present research was to examine the effect of adding a common delay to outcomes in a binary choice, social discounting procedure. Based on the premise that both social and temporal distances are dimensions of psychological distance, we hypothesized that social discounting should decrease as a function of delay to the outcomes. Across two within-subject experiments, participants indicated preference between a hypothetical money reward for the self or for someone else. The outcomes were associated with no, short, and long delays. Both studies confirmed our hypothesis that adding any delay to the receipt of outcomes decreases social discounting, though no significant differences were observed between short and long delays. These results are discussed in the context of some existing literature on altruism.