Fixed Action Patterns Research Articles

A Large-Scale Behavioral Screen to Identify Neurons Controlling Motor Programs in the Drosophila Brain

Drosophila is increasingly used for understanding the neural basis of behavior through genetically targeted manipulation of specific neurons. The primary approach in this regard has relied on the suppression of neuronal activity. Here, we report the results of a novel approach to find and characterize neural circuits by expressing neuronal activators to stimulate subsets of neurons to induce behavior. Classical electrophysiological studies demonstrated that stimulation of command neurons could activate neural circuits to trigger fixed action patterns. Our method was designed to find such command neurons for diverse behaviors by screening flies in which random subsets of brain cells were activated. We took advantage of the large collection of Gal4 lines from the NP project and crossed 835 Gal4 strains with relatively limited Gal4 expression in the brain to flies carrying a UAS transgene encoding TRPM8, a cold-sensitive ion channel. Low temperatures opened the TRPM8 channel in Gal4-expressing cells, leading to their excitation, and in many cases induced overt behavioral changes in adult flies. Paralysis was reproducibly observed in the progeny of crosses with 84 lines, whereas more specific behaviors were induced with 24 other lines. Stimulation performed using the heat-activated channel, TrpA1, resulted in clearer and more robust behaviors, including flight, feeding, and egg-laying. Through follow-up studies starting from this screen, we expect to find key components of the neural circuits underlying specific behaviors, thus providing a new avenue for their functional analysis.

What are Emotional States, and Why Do We Have Them?

An approach to emotion is described in which emotions are defined as states elicited by instrumental reinforcers, that is, by stimuli that are the goals for action. This leads to a theory of the evolutionary adaptive value of emotions, which is that different genes specify different goals in their own self-interest, and any actions can then be learned and performed by instrumental learning to obtain the goals. The brain mechanisms for emotion in brain regions such as the orbitofrontal and anterior cingulate cortex provide a foundation for understanding the neural basis of emotion. Classically conditioned effects may modulate the actions initiated by this system. In addition to this instrumental learning system, some stimuli may elicit responses, for example approach, withdrawal, or fixed action patterns, but intervening states are not required for this type of adaptive response. In addition, a rational thought system involved in multistep planning can allow gene-specified goals to be deferred or avoided in order to achieve longer-term types of goals that may be more advantageous to the individual than to the genes.

An invisible sign stimulus

An object that includes occluded parts is sometimes perceived as a complete image and this phenomenon is known as amodal or visual completion. A sign stimulus is a minimum set of information that elicits a behavior, but this notion raises questions about whether animals ever engage in the behavior when they cannot see the occluded sign stimulus, but they can visually complete it. Male Bengalese finches engage in courtship behavior toward video images of female finches. We conducted three experiments with Bengalese finches to show both sign stimuli and visual completion function in an ecological context. We used three types of visual images recorded from female finches as stimuli: the head, the body, and the whole. Results showed that male Bengalese finches showed courtship behavior toward the head-occluded stimuli whereas they did not toward the headless body image. The results imply that the males completed this occluded sign stimulus through the process of visual completion. Exposure to a sign stimulus combined with the process of visual completion may operate cooperatively to facilitate adaptive responses under conditions of limited information.

Does a nonprimate mammal, the Northern tree shrew (Tupaia belangeri), exhibit paw preference in two forms of a grasping task?

Tree shrews represent a relevant model to study the evolution of primate manual laterality as they are phylogenetically close to primates, they are able to grasp despite having a nonopposable thumb, and they possess a well-developed visual system. In this study, we examined the paw laterality and grasping success rate of 30 Tupaia belangeri (15 males, 15 females) in 2 forced-food grasping tasks (i.e., in a forced-food grasping experiment, the animal has to use paws instead of mouth for food retrieval). We also attempted to determine whether paw usage would be affected by the availability of visual cues using both a visual task (transparent tube) and a nonvisual task (identical but opaque tube). In both tasks, tree shrews showed paw preferences at an individual but not at a population level. Paw laterality (direction and strength) did not differ between tasks. Moreover, in the specific task that we used, grasping success rate was not affected by an absence of visual cues, indicating that tree shrews did not rely on visual guidance to direct their grasps in this forced-food grasping experiment. Our findings suggest that, in contrast to primates, paw usage in tree shrews may result from a modification of a fixed motor pattern in which the preferred direction may be learned. This basic motor organization might be a first step in the evolution of manual laterality, which eventually became controlled by vision in the primate lineage.

Neuroanthropology: Olfactory Recognition of the Self / Non-self by the Ancestral MHC: An EEG Study

Humans naturally discriminate between different ethnic group members using visual and other sensory information, including the sense of smell. The genetic diversity of humans serves as biological and social sign stimuli in the neural self/non-self recognition process. As already evident, the major histocompatibility complex (MHC) encodes both an individual self and a shared ancestral self into the body odor. In a pilot study, we examine the brain-evoked responses to ancestral encoded body odors by conducting an olfactory EEG recording of four different groups: Germans, Taiwanese, Koreans, and Chinese. Results show that humans recognize in-group members as familiar and they exhibit practically identical rhythmic EEG patterns to one’s own odor as it pertains to the ancestral in-group odors. This reveals their genetic relatedness. It is hereby demonstrated that shared ethnicity results in behaviors (both group-protective and pro-social) that are important to intra- and inter-group dynamics.

Neonatal Aspects: Is There Continuity?

ABSTRACT During the 9 months between conception and birth, the fetal brain is transformed from instructions in genes to a complex, highly differentiated organ. The human central nervous system (CNS) changes from a microscopic band of embryonic neuroblasts to a 350 gm mass with more than 109 interconnected highly differentiated neurons in the cortex alone. How this extraordinary growth results in sensomotor, cognitive, affective and behavioral development is still unexplored. The development of voluntary, cognitive and purposive activity from fetal to neonatal period is to analyze the developmental transformations of the brain expressed by development of movement patterns from prenatal through postnatal period. As the development of the brain is unique and continuing process throughout the gestation and after birth, it is expected that there is also continuity of fetal to neonatal movements which are the best functional indicator of developmental processes of the brain. Concerning the complexity, voluntary control and stereotype, there are at least four groups of movements: Reflexes, fixed action patterns, rhythmic motor patterns, and directed movements. Substantial indications suggest that spontaneous activity is a more sensitive indicator of brain dysfunction than reactivity to sensory stimuli in reflex testing. It was proved that assessment of general movements in high-risk newborns has significantly higher predictive value for later neurological development than neurological examination. Nutritional stress at critical times during fetal development can have persistent and potentially irreversible effects on organ function. Impaired intrauterine growth and development may antecede insufficient postnatal growth. Thus, it may be a marker of impaired central nervous system integrity because of adverse intrauterine conditions. Unfavorable intrauterine environment can affect adversely fetal growth. There is an association between postnatal growth and neurodevelopmental outcome. Concerning the continuity from fetus to neonate in terms of neurobehavior, it could be concluded that fetus and neonate are the same persons in different environment. While in the womb, fetus is protected from the gravity which is not so important for its neurodevelopment, postnatally the neonate is exposed to the gravity during the labor and from the first moments of autonomous life. Development of motor control is highly dependent on antigravity forces enabling erect posture of infant or young child. These environmental differences should be kept on mind during prenatal as well as postnatal assessment. How to cite this article Stanojevic M. Neonatal Aspects: Is There Continuity? Donald School J Ultrasound Obstet Gynecol 2012;6(2):189-196.

Fish cognition and behavior

The field of animal cognition is the modern approach to understanding the mental capabilities of animals. The theories are largely an extension of early comparative psychology with a strong influence of behavioural ecology and ethology. Cognition has been variously defined in the literature. Some researchers confine cognition to higher order mental functions including awareness, reasoning and consciousness. However, a more general definition of cognition also includes perception, attention, memory formation and executive functions related to information processing such as learning and problem solving. The study of animal cognition has been largely confined to birds and mammals, particularly non-human primates. This bias in the literature is in part due to the approach taken in the 1950s when cognitive psychologists began to compare known human mental processes with other closely related species. This bias was reinforced by an underlying misconception that learning played little or no role in the development of behaviour in reptiles and fishes. Throughout scientific history fishes have largely been viewed as automatons. Their behaviour was thought to be almost exclusively controlled by unlearned predispositions. Ethologists characterised their behaviour as a series of fixed action patterns released on exposure to appropriate environmental cues (sign stimuli). Whilst there is no doubt that fishes are the most ancient form of vertebrates, they are only ‘primitive’ in the sense that they have been on earth for in excess of 500 million years and that all other vertebrates evolved from some common fish-like ancestor (around 360 million years ago). However, it is important to note that fishes have not been stuck in an evolutionary quagmire during this time. Their form and function have not remained stagnant over the ages. On the contrary, within this time frame they have diversified immensely to the point where there are more species of fish than all other vertebrates combined (currently over 32,000 described species) occupying nearly every imaginable aquatic niche. The erroneous view that both behavioural and neural sophistications are associated in a linear progression from fishes through reptiles and birds to mammals is largely due to a heady mix of outdated and unscientific thinking. Aristotle’s concept of Scala naturae

Intentionality and “free-will” from a neurodevelopmental perspective

The nature of free-will as a subset of intentionality and probabilistic and deterministic function is explored with the indications being that human behavior is highly predictable which in turn, should compromise the notion of free-will. Data supports the notion that age relates to the ability to progressively effectively establish goals performed by fixed action patterns and that these FAPs produce outcomes that in turn modify choices (free-will) for which FAPs need to be employed. Early goals require behaviors that require greater automation in terms of FAPs that lead to goals being achieved or not; if not, then one can change behavior and that in turn is free-will. Goals change with age based on experience which is similar to the way in which movement functions. We hypothesize that human prefrontal cortex development was a natural expansion of the evolutionarily earlier developed areas of the frontal lobe and that goal-directed movements and behavior, including choice and free-will, provided for an expansion of those areas. The same regions of the human central nervous system that were already employed for better control, coordination, and timing of movements, expanded in parallel with the frontal cortex. The initial focus of the frontal lobes was the control of motor activity, but as the movements became more goal-directed, greater cognitive control over movement was necessitated leading to voluntary control of FAPs or free-will. The paper reviews the neurobiology, neurohistology, and electrophysiology of brain connectivities developmentally, along with the development of those brain functions linked to decision-making from a developmental viewpoint. The paper reviews the neurological development of the frontal lobes and inter-regional brain connectivities in the context of optimization of communication systems within the brain and nervous system and its relation to free-will.

Age-Related Change in Phototaxis by Cercariae of Echinostoma caproni (Digenea: Echinostomatidae)

Cercarial dispersal is the result of fixed action patterns in response to reliable environmental cues. We tested the effect of age on the preference of Echinostoma caproni cercariae for light or dark. Individual cercariae were isolated within 10 min of release from Biomphalaria glabrata and placed in a Carolina™ Deep-Well Slide. Half of the slide (top and bottom) was covered with electrical tape to exclude light. The entire chamber of the slide was observed on low power of a dissecting microscope so the cercaria was readily visible whenever it was in the lighted portion of the slide. The amount of time a cercaria spent in the light and the number of times it crossed from light to dark during a 5-min period at 0, 1, 2, and 4 hr were determined (n = 20). The mean amount of time cercariae spent in the light declined significantly from immediately after release (127.7 sec) as compared to 1 hr (68.4 sec), 2 hr (51.6 sec), and 4 hr (10.6 sec) postemergence. The same pattern was seen in the average number of times cercariae crossed from light to dark in a 5-min period: 10.7, 7.2, 6.95, and 1.5, respectively. Cercariae showed no preference for light or dark immediately upon release (P = 0.119), nor was there a correlation between the amount of time spent in the light and the number of crossings at this time period. Cercariae spent a significantly greater amount of time in the dark with age (1–4 hr), and the number of crossings at each of these time periods was highly correlated with the time spent in the light. These findings suggest that light is not an important cue for E. caproni cercariae immediately upon release; however, they develop a strong preference for darker habitats, or an aversion to light, as they age.

No evidence of contagious yawning in the red-footed tortoise Geochelone carbonaria

Abstract Three hypotheses have attempted to explain the phenomenon of contagious yawning. It has been hypothesized that it is a fixed action pattern for which the releasing stimulus is the observation of another yawn, that it is the result of non-conscious mimicry emerging through close links between perception and action or that it is the result of empathy, involving the ability to engage in mental state attribution. This set of experiments sought to distinguish between these hypotheses by examining contagious yawning in a species that is unlikely to show nonconscious mimicry and empathy but does respond to social stimuli: the red-footed tortoise Geochelone carbonaria. A demonstrator tortoise was conditioned to yawn when presented with a red square-shaped stimulus. Observer tortoises were exposed to three conditions: observation of conditioned yawn, non demonstration control, and stimulus only control. We measured the number of yawns for each observer animal in each condition. There was no difference between conditions. Experiment 2 therefore increased the number of conditioned yawns presented. Again, there was no significant difference between conditions. It seemed plausible that the tortoises did not view the conditioned yawn as a real yawn and therefore a final experiment was run using video recorded stimuli. The observer tortoises were presented with three conditions: real yawn, conditioned yawns and empty background. Again there was no significant difference between conditions. We therefore conclude that the red-footed tortoise does not yawn in response to observing a conspecific yawn. This suggests that contagious yawning is not the result of a fixed action pattern but may involve more complex social processes.

Behavioral responses of European sea bass ( Dicentrarchus labrax) larvae and Artemia sp. exposed to constant light or darkness vs. light/dark cycles of white, red or blue wavelengths

The performance and survival of fish larvae are strongly influenced by their surrounding photic environment. The aim of this study was to investigate the effect of light characteristics (spectrum and photoperiod) on the feeding and locomotor behaviors of European sea bass larvae and its prey ( Artemia sp.). To this end, constant light (LL), constant darkness (DD) and 12:12 h LD cycles of red, blue or white LED lights were applied from 1 to 30 days post-hatching. The Modal Action Patterns (swimming duration, orientation, capture, miss and pass frequencies) of larvae and Artemia distribution in the tank were video recorded and analyzed using newly developed tracking software. The results showed that under LD B the phototactic response of sea bass larvae led to a significatively homogeneous distribution in the tanks and aquaria, while under LD W and LL the highest larvae density (52%) was seen on the tank walls. LD B and LD W resulted in longer swimming duration and earlier weaning. Larvae exposed to darkness and red light showed the lowest swimming and feeding activity, and a higher aggregation tendency of both fish larvae and the live prey. White light exposure resulted in a strong phototactic response from fish larvae and Artemia, which consisted of a tendency to congregate at the corners or close to the walls of the tank/aquaria. Artemia hatching rate under blue light was highest (56.5 ± 2.9%) in contrast with red light (26.3 ± 1.4%) and total darkness (27.9 ± 3.9). These results showed that the relationship between the behavioral responses of sea bass larvae and Artemia is strongly affected by lighting conditions, which has both basic and practical implications for understanding their behavioral ecology and for improving culture protocols.

Amusing ourselves to death? Superstimuli and the evolutionary social sciences

Some evolutionary psychologists claim that humans are good at creating superstimuli, and that many pleasure technologies are detrimental to our reproductive fitness. Most of the evolutionary psychology literature makes use of some version of Lorenz and Tinbergen's largely embryonic conceptual framework to make sense of supernormal stimulation and bias exploitation in humans. However, the early ethological concept “superstimulus” was intimately connected to other erstwhile core ethological notions, such as the innate releasing mechanism, sign stimuli and the fixed action pattern, notions that nowadays have, for the most part, been discarded by ethologists. The purpose of this paper is twofold. First, we will reconnect the discussion of superstimuli in humans with more recent theoretical ethological literature on stimulus selection and supernormal stimulation. This will allow for a re-conceptualization of evolutionary psychology's formulation of (supernormal) stimulus selection in terms of domain-specificity and modularity. Second, we will argue that bias exploitation in a cultural species differs substantially from bias exploitation in non-cultural animals. We will explore several of those differences, and explicate why they put important constraints on the use of the superstimulus concept in the evolutionary social sciences.

Source levels of echolocation signals vary in correlation with wingbeat cycle in landing big brown bats (Eptesicus fuscus)

Recordings of the echolocation signals of landing big brown bats with a two-dimensional 16-microphone array revealed that the source level reduction of 7 dB per halving of distance is superimposed by a variation of up to 12 dB within single call groups emitted during the approach. This variation correlates with the wingbeat cycle. The timing of call emission correlates with call group size. First pulses of groups containing many calls are emitted earlier than first calls in groups with fewer calls or single calls. This suggests that the emission of pulse groups follows a fixed motor pattern where the information gained from the preceding pulse group determines how many calls will be emitted in the next group. Single calls and call groups are centred at the middle of the upstroke. Expiration is indicated by call emission. The pause between groups is centred at the middle of the downstroke and indicates inspiration. The hypothesis that the source level variation could be caused by changes in the subglottic pressure due to the contraction of the major flight muscles is discussed.

Analyse der auslösenden Reizsituationen für die Anschwimm-, Eindring- und Fluchtreaktion junger Hemihaplochromis multicolor (Cichlidae)

To investigate the selective response to stimulation the stimuli effecting release of three different reactions in young Hemihaplochromis multicolor were analysed: approach, contact making, and escape. Responses to grey, coloured, and patterned dummies were compared in double-choice experiments. These experiments show that pattern shape has no meaning but that the size of the dark area of the pattern and the contrast in brightness between the dark area and that surrounding it are important sign stimuli for each of the three reactions. Therefore, selective responsiveness does not appear to result from heterogeneous summation but from the quantity of each stimulus parameter and the special combination of sign stimuli.

Half a Century on the Concepts of Innateness and Instinct: Survey, Synthesis and Philosophical Implications

This paper first surveys and criticizes the various attempts by students of behavior to define the concept of innateness. A new definition, based on these (especially that of Lorenz), and reformulable in terms of now classic notions in genetics and evolutionary theory, is then offered: A trait is innate if, and only if, it is canalized and such that any phenotype in the reaction norm which does not possess the trait is a morphose — hence maladapted — and such that all modifications do possess the trait. A narrower definition of innateness, based on the first definition, is suggested to delimit what is called “the Lorenzian notions of innateness” and to provide a framework within which to discuss the notion of a fixed action pattern. Evidence, from the recent literature, is adduced to show the existence and importance of such patterns — even for human behavior. In the final section, it is argued that a paradigm shift has indeed occurred and that although this scientific revolution has been accompanied by many of the irrational elements noted by the Kuhnian school of philosophy of science, the shift has in fact been made upon perfectly rational grounds. That this should be true of so controversial a field is strong evidence against the irrationalist thesis in philosophy of science.

Tits use amodal completion in predator recognition: a field experiment

Amodal completion enables an animal to perceive partly concealed objects as an entirety, and to interact with them appropriately. Several studies, based upon either operant conditioning or filial imprinting techniques, have shown that various animals (both mammals and birds) can perform amodal completion. Before this study, the use of amodal completion by untrained animals in the recognition of objects had not been considered. Using two feeders, we observed in a field experiment the reaction of tits to the torso of a sparrowhawk (partly occluded or an 'amputated' dummy) in two different treatments (sparrowhawk torso vs. complete dummy pigeon; and torso vs. complete dummy sparrowhawk). It is clear that the birds considered the two torso variants as predators and kept away from both of them when the second feeder offered a 'pigeon' instead. On the other hand, when a 'complete sparrowhawk' was present on the second feeder, the number of visits to the occluded torso remained low; while the number of visits to the amputated one increased threefold. Birds risked perching near what was clearly an amputated torso; while the fear of a "hiding" (occluded) torso remained unchanged, when the second feeder did not provide a safe alternative. Such discrimination between torsos requires the ability for amodal completion. Our results demonstrate that in their recognition process, the birds not only use simple sign stimuli, but also complex cognitive functions.

Whatever Happened to Ethology? The Case for the Fixed Action Pattern in Psychology.

In the 1950s ethology entered British psychology as an antidote to learning theory, as a re-incarnation of instinct theory, and as a paradigm for human attachment processes. In the 1970s it became simply the study of “natural” behaviour in an evolutionary framework and was replaced by socio-biology and evolutionary psychology, in which species comparisons and evolutionary relationships enable adaptational (i.e. functional) explanations of human behaviours. Ethological theory had been abandoned. Yet the concept of the fixed action pattern (FAP) is the behavioural equivalent of the species in biology. Both identify constancies in variety. Together with the concepts sign stimulus, innate releaser, specific action potential (SAP), displacement activity, appetitive and consummatory behaviour, complex behaviour can be understood as interacting elements for which there should be neuro-hormonal and genetic systems (i.e. mechanisms). An analysis of the nature of emotion using ethological theory is presented as an example. Neuro-ethological work began in the 1960s with brain lesions and stimulation, and continues with modern technologies. For example, syllables in bird song (FAPs?) are correlated with activity in a striatal brain region (HVc) and a gene (FoxP2) – a gene that affects speech. An outline of the ethology and neuro-ethology of speech is presented.

Virgin queen execution in the stingless beeMelipona beecheii: The sign stimulus for worker attacks

Stingless bees produce considerable numbers of virgin queens over the year, most of which are superfluous and get executed by the workers. Nothing is known yet about the sign stimulus that releases the worker attacks. In the present study we investigated the queen execution process in Melipona beecheii and found in both behavioral observations and experiments with caged living virgin queens that workers are not attracted to them from a distance by means of volatile chemicals. Furthermore, worker aggression, which is obvious towards virgin queens that run through the nest excitedly with their abdomen enlarged and the wings beating, was lowered to almost zero when we made the queens behavior-less by experimentally killing them. Our results clearly show that the sign stimulus for releasing the execution behavior in Melipona beecheii workers is not a chemical stimulus but the virgin queens' conspicuous behavior, which, we hypothesize, could act as a direct measure of their fitness.

Characterization of the Decision Network for Wing Expansion inDrosophilaUsing Targeted Expression of the TRPM8 Channel

After emergence, adult flies and other insects select a suitable perch and expand their wings. Wing expansion is governed by the hormone bursicon and can be delayed under adverse environmental conditions. How environmental factors delay bursicon release and alter perch selection and expansion behaviors has not been investigated in detail. Here we provide evidence that in Drosophila the motor programs underlying perch selection and wing expansion have different environmental dependencies. Using physical manipulations, we demonstrate that the decision to perch is based primarily on environmental valuations and is incrementally delayed under conditions of increasing perturbation and confinement. In contrast, the all-or-none motor patterns underlying wing expansion are relatively invariant in length regardless of environmental conditions. Using a novel technique for targeted activation of neurons, we show that the highly stereotyped wing expansion motor patterns can be initiated by stimulation of N(CCAP), a small network of central neurons that regulates the release of bursicon. Activation of this network using the cold-sensitive rat TRPM8 channel is sufficient to trigger all essential behavioral and somatic processes required for wing expansion. The delay of wing expansion under adverse circumstances thus couples an environmentally sensitive decision network to a command-like network that initiates a fixed action pattern. Because N(CCAP) mediates environmentally insensitive ecdysis-related behaviors in Drosophila development before adult emergence, the study of wing expansion promises insights not only into how networks mediate behavioral choices, but also into how decision networks develop.

EXCITATION AND HABITUATION OF CRAYFISH ESCAPE

This paper describes one of the first attempts to analyze the synaptic basis for the release of an animal's fixed action pattern, which controls a behavior such as the crayfish escape response. It is also one of the early demonstrations that the neural mechanisms of a simple form of learning, known