Contributions of testosterone and territory ownership to sexually-motivated behaviors and mRNA expression in the medial preoptic area of male European starlings

Animals integrate social information with their internal endocrine state to control the timing of behavior, but how these signals are integrated in the brain is not understood. The medial preoptic area (mPOA) may play an integrative role in the control of courtship behavior, as it receives projections from multiple sensory systems, and is central to the hormonal control of courtship behavior across vertebrates. Additionally, data from many species implicate opioid and dopaminergic systems in the mPOA in the control of male courtship behavior. We used European starlings to test the hypothesis that testosterone (T) and social status (in the form of territory possession) interact to control the timing of courtship behavior by modulating steroid hormone-, opioid- and dopaminergic-related gene expression in the mPOA. We found that only males given both T and a nesting territory produced high rates of courtship behavior in response to a female. T treatment altered patterns of gene expression in the mPOA by increasing androgen receptor, aromatase, mu-opioid receptor and preproenkephalin mRNA and decreasing tyrosine hydroxylase mRNA expression. Territory possession did not alter mRNA expression in the mPOA, despite the finding that only birds with both T and a nesting territory produced courtship behavior. We propose that T prepares the mPOA to respond to the presence of a female with high rates of courtship song by altering gene expression, but that activity in the mPOA is under a continuous (i.e. tonic) inhibition until a male starling obtains a nesting territory.     

Behavior of Early Warnings near the Critical Temperature in the Two-Dimensional Ising Model

Among the properties that are common to complex systems, the presence of critical thresholds in the dynamics of the system is one of the most important. Recently, there has been interest in the universalities that occur in the behavior of systems near critical points. These universal properties make it possible to estimate how far a system is from a critical threshold. Several early-warning signals have been reported in time series representing systems near catastrophic shifts. The proper understanding of these early-warnings may allow the prediction and perhaps control of these dramatic shifts in a wide variety of systems. In this paper we analyze this universal behavior for a system that is a paradigm of phase transitions, the Ising model. We study the behavior of the early-warning signals and the way the temporal correlations of the system increase when the system is near the critical point.     

Signal acquisition and analysis for cortical control of neuroprosthetics

Work in cortically controlled neuroprosthetic systems has concentrated on decoding natural behaviors from neural activity, with the idea that if the behavior could be fully decoded it could be duplicated using an artificial system. Initial estimates from this approach suggested that a high-fidelity signal comprised of many hundreds of neurons would be required to control a neuroprosthetic system successfully. However, recent studies are showing hints that these systems can be controlled effectively using only a few tens of neurons. Attempting to decode the pre-existing relationship between neural activity and natural behavior is not nearly as important as choosing a decoding scheme that can be more readily deployed and trained to generate the desired actions of the artificial system. These artificial systems need not resemble or behave similarly to any natural biological system. Effective matching of discrete and continuous neural command signals to appropriately configured device functions will enable effective control of both natural and abstract artificial systems using compatible thought processes.     

The Evolution of Neuroanatomical Substrates of Reproductive Behavior: Sex Steroid and LHRH-Specific Pathways Including the Terminal Nerve

Fairly recent anatomical methods have made possible the mappingof neurobehavioral systems involving two types of reproductivehormones, gonadal steroids and the peptide luteinizing hormonereleasing hormone (LHRH). Brain sites of steroid uptake aredetected using autoradiography; LHRH is localized in cells andfibers using immunocytochemical procedures. Both hormone typesare known to strongly influence sex behavior and it can reasonablybe assumed that these effects are mediated in large part viasystems identified using the anatomical procedures. Analysisof the comparative anatomy of these systems should thereforeprovide information useful in the construction of models concerningthe evolution of neurohormonal control of reproductive behavior.The results of such a study are reported. Sex steroid and LHRHsystems in cyclostomes, teleosts, amphibians, reptiles, birdsand mammals are considered in detail. A synthesis of this informationhas led to the following ideas. Androgenic control of male reproductivesystems has evolved in a number of nonhomologous motor systemscontrolling male reproductive behavior. Sex steroid and LHRHsystems may interact at several different levels of the neuraxisbut the most obvious overlap of the systems occurs in the septaland POA areas. The latter especially is a fairly constant andperhaps primitive feature. LHRH secretion into the systemiccirculation was most likely the earliest means for LHRH modulationof both pituitary function and neural systems controlling reproductivebehavior.Pathways for more direct delivery of LHRH to pituitarycells and brain nuclei probably developed in the early gnathostomes.The terminal nerve appears to be a rather conservative LHRH-containingpathway connecting olfactory systems with septal-preoptic nuclei.A function in pheromonal control of sex behavior is suggested.The general distribution of steroid concentrating cells andLHRH pathways in tetrapods seems to be rather constant. Absenceof the systems in neocortical areas and their homologs is conspicuous.     

Estrogen and endogenous opioids regulate CCK in reproductive circuits.

This review focuses on the interaction of estrogen with the cholecystokinin (CCK) and endogenous opioid peptide systems in the medial preoptic nucleus, and how these interactions result in alterations of a stereotypic female reproductive behavior--lordosis. The medial preoptic nucleus is an integral part of a circuit controlling lordosis that extends from the limbic system through the hypothalamus. Estrogen alters the integration of sensory information in the circuit that results in the display of sexually receptive behavior. Estrogen determines the activity of CCK and endogenous opioid peptide systems through regulation of expression, release and interaction with specific receptors. Studies of each system individually have indicated that they are pivotal to the expression of lordosis. Recent studies demonstrate an estrogen-dependent interaction between endogenous opioid and CCK systems that control reproductive behavior.     

Oscillations in Controlled Biochemical Systems

Stability analysis of equations describing certain biochemical control mechanisms involving negative feedback suggests that limit cycle behavior might be possible if the control system involves a sufficient number of intermediate chemical steps. For the example considered in this paper, digital simulation of the non-linear control system illustrates that limit cycle behavior actually arises for a sixth-order system. On the other hand, the corresponding fourth- and fifth-order systems are asymptotically stable.     

Dynamic control in metabolic engineering: Theories,tools, and applications

Metabolic engineering has allowed the production of a diverse number of valuable chemicals using microbial organisms. Many biological challenges for improving bio-production exist which limit performance and slow the commercialization of metabolically engineered systems. Dynamic metabolic engineering is a rapidly developing field that seeks to address these challenges through the design of genetically encoded metabolic control systems which allow cells to autonomously adjust their flux in response to their external and internal metabolic state. This review first discusses theoretical works which provide mechanistic insights and design choices for dynamic control systems including two-stage, continuous, and population behavior control strategies. Next, we summarize molecular mechanisms for various sensors and actuators which enable dynamic metabolic control in microbial systems. Finally, important applications of dynamic control to the production of several metabolite products are highlighted, including fatty acids, aromatics, and terpene compounds. Altogether, this review provides a comprehensive overview of the progress, advances, and prospects in the design of dynamic control systems for improved titer, rate, and yield metrics in metabolic engineering.     

Capital punishment in tribal Montenegro: Implications for law,biology, and theory of social control

Clan execution among feuding tribal societies is proposed as a significant precursor to modern law. This form of ostracism is examined with respect to its social control functions and the indigenous assumptions about behavior, genes and demography that guide the behavior. It is suggested that there are very close parallels between modern legal systems and nonliterate ones based on “self-help”, when capital punishment is used by normal decision groups in the name of “national security”.     

Adolescent cognitive control and reward processing: Implications for risk taking and substance use

Adolescence is a unique, transitional period of human development. Once hallmark of this period is progressive improvements (relative to children) in cognitive control, core mental abilities enabling the ‘top-down’, endogenous control over behavior. However, as adolescents transition to more mature (adult) levels of functioning, limitations still exist in the ability to consistently and flexibly exert cognitive control across various contexts into the early twenties. Adolescence is also marked by peaks in sensation, novelty, and reward seeking behaviors thought to stem from normative increases in responsiveness in limbic and paralimbic brain structures, beginning around the onset of puberty. Asynchronous maturation in these systems during the adolescent period likely contributes to immature decision-making, strongly influenced by ‘bottom-up’ reward processes, and may help explain noted increases in risk taking behavior during adolescence. In this paper, structural and functional maturation in brain systems supporting reward and cognitive control processing are reviewed as a means to better understand risk taking. Particular emphasis is placed on adolescents' experimentation with drugs as a specific example of a risky behavior.     

Fruit fly behavior in response to chemosensory signals

An important question in contemporary sensory neuroscience is how animals perceive their environment and make appropriate behavioral choices based on chemical perceptions. The fruit fly Drosophila melanogaster exhibits robust tastant and odor-evoked behaviors. Understanding how the gustatory and olfactory systems support the perception of these contact and volatile chemicals and translate them into appropriate attraction or avoidance behaviors has made an unprecedented contribution to our knowledge of the organization of chemosensory systems. In this review, I begin by describing the receptors and signaling mechanisms of the Drosophila gustatory and olfactory systems and then highlight their involvement in the control of simple and complex behaviors. The topics addressed include feeding behavior, learning and memory, navigation behavior, neuropeptide modulation of chemosensory behavior, and I conclude with a discussion of recent work that provides insight into pheromone signaling pathways.     

Sensorimotor control of navigation in arthropod and artificial systems

Arthropods exhibit highly efficient solutions to sensorimotor navigation problems. They thus provide a source of inspiration and ideas to robotics researchers. At the same time, attempting to re-engineer these mechanisms in robot hardware and software provides useful insights into how the natural systems might work. This paper reviews three examples of arthropod sensorimotor control systems that have been implemented and tested on robots. First we discuss visual control mechanisms of flies, such as the optomotor reflex and collision avoidance, that have been replicated in analog VLSI (very large scale integration) hardware and used to produce corrective behavior in robot vehicles. Then, we present a robot model of auditory localization in the cricket; and discuss integration of this behavior with the optomotor behavior previously described. Finally we present a model of olfactory search in the moth, which makes use of several sensory cues, and has also been tested using robot hardware. We discuss some of the similarities and differences of the solutions obtained.     

Optimal foraging and predator-prey dynamics, II.

In this paper we consider one-predator-two-prey population dynamics described by a control system. We study and compare conditions for permanence of the system for three types of predator feeding behaviors: (i) specialized feeding on the more profitable prey type, (ii) generalized feeding on both prey types, and (iii) optimal foraging behavior. We show that the region of parameter space leading to permanence for optimal foraging behavior is smaller than that for specialized behavior, but larger than that for generalized behavior. This suggests that optimal foraging behavior of predators may promote coexistence in predator-prey systems. We also study the effects of the above three feeding behaviors on apparent competition between the two prey types.     

On the applicability of the decentralized control mechanism extracted from the true slime mold: a robotic case study with a serpentine robot

A systematic method for an autonomous decentralized control system is still lacking, despite its appealing concept. In order to alleviate this, we focused on the amoeboid locomotion of the true slime mold, and extracted a design scheme for the decentralized control mechanism that leads to adaptive behavior for the entire system, based on the so-called discrepancy function. In this paper, we intensively investigate the universality of this design scheme by applying it to a different type of locomotion based on a 'synthetic approach'. As a first step, we implement this design scheme to the control of a real physical two-dimensional serpentine robot that exhibits slithering locomotion. The experimental results show that the robot exhibits adaptive behavior and responds to the environmental changes; it is also robust against malfunctions of the body segments due to the local sensory feedback control that is based on the discrepancy function. We expect the results to shed new light on the methodology of autonomous decentralized control systems.     

Bifurcation theory explains waveform variability in a congenital eye movement disorder

In dynamical systems, configurations that permit flexible control are also prone to undesirable behavior. We study a bilateral model of the oculomotor pre-motor network that conforms with the neuroanatomical constraint that brainstem neurons project to cerebellar Purkinje cells on both sides, but Purkinje cells project back to brainstem neurons on the same side only. Bifurcation analysis reveals that this network asymmetry enables flexible control by the cerebellum of brainstem network dynamics, but small changes in connection pattern or strength lead to behavior that is unstable, oscillatory, or both. The model produces the full range of waveform types associated with the hereditary eye movement disorder know as congenital nystagmus, and is consistent with findings linking the disorder with abnormal connectivity or limited plasticity in the cerebellum.     

A theoretical framework for CNS arousal

Rapid changes of state in central nervous systems (CNS), as required following stimuli that must arouse the CNS from a quiescent state in order to activate a behavioral response, constitute a particularly appropriate application of non-linear dynamics. Chaotic dynamics would provide tremendous amplification of neuronal activity needed for CNS arousal, sensitively dependent on the initial state of the CNS. This theoretical approach is attractive because it supposes dynamics that are deterministic and it links the elegant mathematics of chaos to the conception of a fundamental property of the CNS. However, a living system must be able to exit from chaotic dynamics in order to avoid widely divergent, biologically impossible outcomes. We hypothesize that, analogous to phase transitions in a liquid crystal, CNS arousal systems, having 'woken up the brain' to activate behavior, go through a phase transition and emerge under the control of orderly movement control systems.     

Slugs and snails and opiate tales: opioids and feeding behavior in invertebrates

There is accumulating evidence that opioid systems are involved in the regulation of fundamental behavioral and physiological processes in invertebrates. Feeding is a basic physiological function that is essential for maintaining homeostasis. Results of studies examining the feeding responses of molluscs and arthropods treated with various opiate agonists and antagonists indicate that delta, kappa, mu, and possibly sigma opioid systems differentially and selectively mediate the components of their natural feeding behavior. Moreover, it appears that at an early evolutionary stage the mu and kappa systems have developed to selectively affect the components of feeding behavior associated with the acquisition and ingestion of food. In addition, evidence suggests that neuropeptides that have been proposed as possible endogenous antagonists of opioid-mediated feeding in mammals may also be involved in the control of feeding in invertebrates. This indicates that there may be an interplay of opioid agonists and antagonists in the regulation of feeding and satiation in invertebrates analogous to that proposed for vertebrates. Moreover, these findings indicate that opioid influences on feeding have been conserved through evolution.     

A facet of the man-child bond: The teeter-totter effect

Evidence is presented for the candidacy of the adult male-child bond as a species-characteristic behavior. The evidence includes (a) catholicity of trait, (b) arbitrariness of trait, (c) adequate number of observations to allow intracultural norms to emerge, (d) substantial incidence of the trait's occurrence, and (e) accurate predictability of facets of the trait. If the candidacy is successful, then the adult male-child bond is hypothesized to be a generalized or thematic behavior pattern that is canalized and biased by (autogenetic and active) processes within the motivation systems—the nervous system and the endocrine system. These systems, in turn, are under partial control of the genetic constellations which are characteristic of the species Homo sapiens.     

Instantaneous flux control analysis for biochemical systems.

Linear sensitivity analysis of steady-state control of enzymic systems has been extended to non-steady states yielding sensitivity coefficients which provide non-intuitive insights into the behavior of the system and the sites of metabolic control, and which are quantitative counterparts to traditional qualitative concepts. Because this information is provided in a readily understood format, these coefficients serve as convenient indices of metabolic control. This treatment was applied to a simple test system, consisting of two enzymes and one non-enzymatic reaction, which exhibits oscillatory behavior. The results indicate that oscillations in the concentrations of the intermediate metabolites are regulated almost exclusively by the second enzyme. Control of the flux through the pathway is apportioned equally among the three reactions during periods of low net flux, but it is due almost exclusively to the second enzyme during periods of high net flux.     

A genetic approach to dissect sexually dimorphic behaviors

It has been known since antiquity that gender-specific behaviors are regulated by the gonads. We now know that testosterone is required for the appropriate display of male patterns of behavior. Estrogen and progesterone, on the other hand, are essential for female typical responses. Research from several groups also indicates that estrogen signaling is required for male typical behaviors. This finding raises the issue of the relative contribution of these two hormonal systems in the control of male typical behavioral displays. In this review we discuss the findings that led to these conclusions and suggest various genetic strategies that may be required to understand the relative roles of testosterone and estrogen signaling in the control of gender-specific behavior.     

Mechanical aspects of legged locomotion control

We review the mechanical components of an approach to motion science that enlists recent progress in neurophysiology, biomechanics, control systems engineering, and non-linear dynamical systems to explore the integration of muscular, skeletal, and neural mechanics that creates effective locomotor behavior. We use rapid arthropod terrestrial locomotion as the model system because of the wealth of experimental data available. With this foundation, we list a set of hypotheses for the control of movement, outline their mathematical underpinning and show how they have inspired the design of the hexapedal robot, RHex.