A computer-based avatar task can differentiate avoidant and non-avoidant coping styles.

Background/Objective: Coping styles play a role in how individuals respond to stress and therapy. One aspect of coping which has been linked to adverse outcomes including anxiety disorders and PTSD is avoidance. However, a tendency to avoid may affect the accuracy of paper and pencil inventories used to identify avoidant temperaments. Previously, we showed that a computer-based task in which an on-screen "avatar" is guided through a series of onscreen events could predict avoidance including behavioral inhibition, harm avoidance, and self-reported PTSD symptoms. Since some coping styles involve avoidance, we extended this work to determine whether scores on the avatar task would also differentiate avoidant and non-avoidant coping styles as measured by the Brief COPE. Methods: One hundred and fifty undergraduates voluntarily completed the avatar task and the Brief COPE. Results: Scores on the avatar task had a significant positive relationship with an aggregate score for the five avoidant coping styles and a significant negative relationship with an aggregate score for the nine non-avoidant coping styles. Conclusions: The effectiveness of the avatar task to differentiate coping styles based on avoidance further validates this task and also shows selectivity to avoidant coping styles as opposed to other non-avoidant coping styles.

Avoidance learning and classical eyeblink conditioning as model systems to explore a learning diathesis model of PTSD.

Experiencing a trauma is necessary, but not sufficient, for the development of post-traumatic stress disorder (PTSD) in that most individuals who experience a trauma do not go on to develop PTSD. This suggests that identifiable vulnerabilities (i.e., diatheses) exist that increase the risk for the development of PTSD. One such factor is the personality temperament of behavioral inhibition (BI). Organisms that exhibit BI were studied in the context of avoidance learning and classical eyeblink conditioning. We present a body of evidence supporting a learning diathesis model in which behaviorally inhibited organisms exhibit enhanced acquisition and resistance to extinction in these tasks. Vulnerable individuals show learning-related enhancements when the learning situation involves some degree of uncertainty. We review the known brain circuitry involved in classical eyeblink conditioning in the context of the learning diathesis model. Finally, the data reviewed here demonstrate the value of studying vulnerability factors in humans and a rodent model using cerebellar-dependent learning tasks for understanding the acquisition and endurance of PTSD symptomatology.

The distressed (Type D) personality factor of social inhibition, but not negative affectivity, enhances eyeblink conditioning.

Recent work has focused on a learning diathesis model in which specific personality factors such as behavioral inhibition (BI) may influence associative learning and in turn increase risk for the development of anxiety disorders. We have found in a series of studies that individuals self-reporting high levels of BI exhibit enhanced acquisition of conditioned eyeblinks. In the study reported here, hypotheses were extended to include distressed (Type D) personality which has been found to be related to BI. Type D personality is measured with the DS-14 scale which includes two subscales measuring negative affectivity (NA) and social inhibition (SI). We hypothesized that SI, which is similar to BI, would result in enhanced acquisition while the effect of NA is unclear. Eighty nine participants completed personality inventories including the Adult Measure of Behavioral Inhibition (AMBI) and DS-14. All participants received 60 acquisition trials with a 500 ms, 1000 Hz, tone CS and a co-terminating 50 ms, 5 psi corneal airpuff US. Participants received either 100% CS-US paired trials or a schedule of partial reinforcement where 50% US alone trials were intermixed into CS-US training. Acquisition of CRs did not differ between the two training protocols. Whereas BI was significantly related to Type D, SI, and NA, only BI and SI individuals exhibited enhanced acquisition of conditioned eyeblinks as compared to non-inhibited individuals. Personality factors now including social inhibition can be used to identify individuals who express enhanced associative learning which lends further support to a learning diathesis model of anxiety disorders.

US alone trials presented during acquisition do not disrupt classical eyeblink conditioning: Empirical and computational findings.

Studies of partial reinforcement in eyeblink conditioning have typically shown slower learning of a CS-US association when paired CS-US trials are interleaved with CS-alone trials. However, recent work has shown that CS-US learning is not slowed by interleaved US-alone trials. This discrepancy is surprising since both partial reinforcement protocols reduce the total number of paired CS-US trials. Previously, Kimble et al. (1955) reported that inserting a block of US-alone trials during CS-US training did not disrupt eyeblink acquisition. Here, we sought to replicate and extend these findings by comparing interleaved vs. blocked US-alone trials during CS-US paired training. Ninety-seven undergraduates volunteered for this experiment for research credit. Participants received 60 acquisition trials, consisting of either 100% CS-US paired trials, 50% US-alone trials intermixed with CS-US paired trials, or a block of 20 US-alone trials inserted between blocks of 20 CS-US trials. We also utilized a previously published computational model of hippocampal and cerebellar learning to test the effects of these US-alone protocols. Both empirical and computational results supported the finding that US-alone trials, either intermixed or inserted as a block of trials, do not disrupt acquisition of conditioned eyeblinks. Possible neural substrates of these US-alone effects are discussed.

Uncertainty of trial timing enhances acquisition of conditioned eyeblinks in anxiety vulnerable individuals.

Recent work has found that behaviorally inhibited (BI) individuals exhibit enhanced eyeblink conditioning in omission and yoked training as well as with schedules of partial reinforcement. We hypothesized that spacing CS-US paired trials over a longer period of time by extending and varying the inter-trial interval (ITI) would facilitate learning. All participants completed the Adult Measure of Behavioural Inhibition (AMBI) and were grouped as behaviorally inhibited (BI) and non-behaviorally inhibited (NI) based on a median split score of 15.5. All participants received 3 US alone trials and 30CS-US paired trials for acquisition training and 20CS alone trials for extinction training in one session. Conditioning stimuli were a 500 ms tone conditioned stimulus (CS) and a 50-ms air puff unconditional stimulus (US). Participants were randomly assigned to receive a short ITI (mean=30+/- 5s), a long ITI (mean=57+/- 5s) or a variable long ITI (mean=57 s, range 25-123 s). No significant ITI effects were observed for acquisition or extinction. Overall, anxiety vulnerable individuals exhibited enhanced conditioned eyeblink responses as compared to non-vulnerable individuals. This enhanced acquisition of CRs was significant in spaced training with a variable long ITI, but not the short or long ITI. There were no significant effects of ITI or BI on extinction. These findings are interpreted based on the idea that uncertainty plays a role in anxiety and can enhance associative learning in anxiety vulnerable individuals.

Behaviorally inhibited individuals demonstrate significantly enhanced conditioned response acquisition under non-optimal learning conditions.

Behavioral inhibition (BI) is an anxiety vulnerability factor associated with hypervigilance to novel stimuli, threat, and ambiguous cues. The progression from anxiety risk to a clinical disorder is unknown, although the acquisition of defensive learning and avoidance may be a critical feature. As the expression of avoidance is also central to anxiety development, the present study examined avoidance acquisition as a function of inhibited temperament using classical eyeblink conditioning. Individuals were classified as behaviorally inhibited (BI) or non-inhibited (NI) based on combined scores from the Adult and Retrospective Measures of Behavioural Inhibition (AMBI and RMBI, respectively). Acquisition was assessed using delay, omission, or yoked conditioning schedules of reinforcement. Omission training was identical to delay, except that the emission of an eyeblink conditioned response (CR) resulted in omission of the unconditioned airpuff stimulus (US) on that trial. Each subject in the yoked group was matched on total BI score to a subject in the omission group, and received the same schedule of CS and US delivery, resulting in a partial reinforcement training schedule. Delay conditioning elicited significantly more CRs compared to the omission and yoked contingencies, the latter two of which did not differ from each other. Thus, acquisition of an avoidance response was not apparent. BI individuals demonstrated enhanced acquisition overall, while partial reinforcement training significantly distinguished between BI and NI groups. Enhanced learning in BI may be a function of an increased defensive learning capacity, or sensitivity to uncertainty. Further work examining the influence of BI on learning acquisition is important for understanding individual differences in disorder etiology in anxiety vulnerable cohorts.

Gate-defined quantum confinement in suspended bilayer graphene.

Quantum-confined devices that manipulate single electrons in graphene are emerging as attractive candidates for nanoelectronics applications. Previous experiments have employed etched graphene nanostructures, but edge and substrate disorder severely limit device functionality. Here we present a technique that builds quantum-confined structures in suspended bilayer graphene with tunnel barriers defined by external electric fields that open a bandgap, thereby eliminating both edge and substrate disorder. We report clean quantum dot formation in two regimes: at zero magnetic field B using the energy gap induced by a perpendicular electric field and at B>0 using the quantum Hall ν=0 gap for confinement. Coulomb blockade oscillations exhibit periodicity consistent with electrostatic simulations based on local top-gate geometry, a direct demonstration of local control over the band structure of graphene. This technology integrates single electron transport with high device quality and access to vibrational modes, enabling broad applications from electromechanical sensors to quantum bits.

Broken-symmetry states in doubly gated suspended bilayer graphene.

The single-particle energy spectra of graphene and its bilayer counterpart exhibit multiple degeneracies that arise through inherent symmetries. Interactions among charge carriers should spontaneously break these symmetries and lead to ordered states that exhibit energy gaps. In the quantum Hall regime, these states are predicted to be ferromagnetic in nature, whereby the system becomes spin polarized, layer polarized, or both. The parabolic dispersion of bilayer graphene makes it susceptible to interaction-induced symmetry breaking even at zero magnetic field. We investigated the underlying order of the various broken-symmetry states in bilayer graphene suspended between top and bottom gate electrodes. We deduced the order parameter of the various quantum Hall ferromagnetic states by controllably breaking the spin and sublattice symmetries. At small carrier density, we identified three distinct broken-symmetry states, one of which is consistent with either spontaneously broken time-reversal symmetry or spontaneously broken rotational symmetry.

Local compressibility measurements of correlated states in suspended bilayer graphene.

Bilayer graphene has attracted considerable interest due to the important role played by many-body effects, particularly at low energies. Here we report local compressibility measurements of a suspended graphene bilayer. We find that the energy gaps at filling factors ν= ± 4 do not vanish at low fields, but instead merge into an incompressible region near the charge neutrality point at zero electric and magnetic field. These results indicate the existence of a zero-field ordered state and are consistent with the formation of either an anomalous quantum Hall state or a nematic phase with broken rotational symmetry. At higher fields, we measure the intrinsic energy gaps of broken-symmetry states at ν=0, ± 1, and ± 2, and find that they scale linearly with magnetic field, yet another manifestation of the strong Coulomb interactions in bilayer graphene.

Using L-systems for modeling source-sink interactions, architecture and physiology of growing trees: the L-PEACH model.

Functional-structural plant models simulate the development of plant structure, taking into account plant physiology and environmental factors. The L-PEACH model is based on the development of peach trees. It demonstrates the usefulness of L-systems in constructing functional-structural models. L-PEACH uses L-systems both to simulate the development of tree structure and to solve differential equations for carbohydrate flow and allocation. New L-system-based algorithms are devised for simulating the behavior of dynamically changing structures made of hundreds of interacting, time-varying, nonlinear components. L-PEACH incorporates a carbon-allocation model driven by source-sink interactions between tree components. Storage and mobilization of carbohydrates during the annual life cycle of a tree are taken into account. Carbohydrate production in the leaves is simulated based on the availability of water and light. Apices, internodes, leaves and fruit grow according to the resulting local carbohydrate supply. L-PEACH outputs an animated three-dimensional visual representation of the growing tree and user-specified statistics that characterize selected stages of plant development. The model is applied to simulate a tree's response to fruit thinning and changes in water stress. L-PEACH may be used to assist in horticultural decision-making processes after being calibrated to specific trees.

Cerebellar substrates for error correction in motor conditioning.

The authors evaluate a mapping of Rescorla and Wagner's (1972) behavioral model of classical conditioning onto the cerebellar substrates for motor reflex learning and illustrate how the limitations of the Rescorla-Wagner model are just as useful as its successes for guiding the development of new psychobiological theories of learning. They postulate that the inhibitory pathway that returns conditioned response information from the cerebellar interpositus nucleus back to the inferior olive is the neural basis for the error correction learning proposed by Rescorla and Wagner (Gluck, Myers, & Thompson, 1994; Thompson, 1986). The authors' cerebellar model expects that behavioral processes described by the Rescorla-Wagner model will be localized within the cerebellum and related brain stem structures, whereas behavioral processes beyond the scope of the Rescorla-Wagner model will depend on extracerebellar structures such as the hippocampus and related cortical regions. Simulations presented here support both implications. Several novel implications of the authors' cerebellar error-correcting model are described including a recent empirical study by Kim, Krupa, and Thompson (1998), who verified that suppressing the putative error correction pathway should interfere with the Kamin (1969) blocking effect, a behavioral manifestation of error correction learning. The authors also discuss the model's implications for understanding the limits of cerebellar contributions to associative learning and how this informs our understanding of hippocampal function in conditioning. This leads to a more integrative view of the neural substrates of conditioning in which the authors' real-time circuit-level model of the cerebellum can be viewed as a generalization of the long-term memory module of Gluck and Myers' (1993) trial-level theory of cerebellar-hippocampal interaction in motor conditioning.

Parallel neural systems for classical conditioning: support from computational modeling.

Classical conditioning has been explained by two main types of theories that postulate different learning mechanisms. Rescorla and Wagner (1972) put forth a theory in which conditioning is based on the ability of the US to drive learning through error correction. Alternatively, Mackintosh (1973) put forth a theory in which the ability of the CS to be associated with the unconditioned stimulus is modulated. We have proposed a reconciliation of these two mechanisms as working in parallel within different neural systems: a cerebellar system for US modulation and a hippocampal system for CS modulation. We developed a computational model of cerebellar function in eyeblink conditioning based on the error correction mechanism of the Rescorla-Wagner rule in which learning-related activity from the cerebellum inhibits the inferior olive, which is the US input pathway to the cerebellum (Gluck et al., 1994). We developed a computational model of the hippocampal region that forms altered representations of conditioned stimuli based on their behavioral outcomes (Gluck & Myers, 1993; Myers et al., 1995). Overall, computational modeling and empirical findings support the idea that, at least in the case of eyeblink conditioning, there may be two different neural systems: the cerebellum which mediates US-based error correction and hippocampus which alters representations of CSs.

Patterns of sympathetic and parasympathetic reactivity in a sample of children and adolescents.

We hypothesized that patterns of sympathetic and parasympathetic reactivity observed in adults would be apparent in a sample of children and adolescents and that these patterns would be consistent across tasks. We explored the relationship between these patterns and psychosocial risk factors for cardiovascular disease. We measured preejection period (PEP) and an index of respiratory sinus arrhythmia (mean successive difference [MSD] statistic) during three reactivity tasks. We classified participants into four groups based on increases or decreases in PEP and MSD. Ninety percent of the sample exhibited the same pattern during at least two of the tasks. PEP and MSD demonstrated consistency, suggesting individual response stereotypy. Exhibiting a consistent pattern of decreased PEP and increased MSD was associated with less child- and parent-reported family conflict. These results are discussed in the context of vagal regulation of environmental demands.

Dissociating entorhinal and hippocampal involvement in latent inhibition.

This study used anatomical cues to suggest a functional dissociation between the roles of the entorhinal cortex and the hippocampus in learning. The authors proposed that the highly convergent inputs to the entorhinal cortex indicate this region may be particularly important for selecting or compressing information. This hypothesis was tested in rabbits (Oryctolagus cunniculus) trained on an associative learning task that is a common index of stimulus selection. In this task, known as latent inhibition, preexposure to a stimulus (such as a tone) leads to slowed learning when the same tone is subsequently paired with an outcome (such as an airpuff to the eye). As hypothesized, rabbits with neurotoxic lesions of the entorhinal cortex failed to show slowed learning following preexposure (no latent inhibition) and learned the association faster than control rabbits. In contrast, hippocampal-lesioned animals showed normal (slowed) learning.

The relationships of resting baroreflex sensitivity, heart rate variability and measures of impulse control in children and adolescents.

The objectives of the present study were to assess: (1) the feasibility of using a non-invasive method to measure baroreflex sensitivity in children and adolescents; (2) the relationships of resting baroreflex sensitivity with resting levels of other cardiovascular variables; and (3) whether baroreflex sensitivity and heart rate variability, two indices of cardiac vagal control at rest, were related to measures of impulse control. Ninety-one Black and White children (ages 8-10) and adolescents (ages 15-17), both female and male, participated in the study. Baroreflex sensitivity, auscultatory blood pressure, EKG-derived heart rate, and the mean successive difference of interbeat intervals were collected during a 10-min rest period. Measures of impulse control came from the Interview for Antisocial Behavior. Baroreflex sensitivity was strongly positively correlated with mean successive difference and negatively correlated with heart rate for all participant groups; baroreflex sensitivity was negatively correlated with diastolic blood pressure, but only for children, males, and Blacks. Increased problems with impulse control was associated with decreased cardiac vagal control, but only in males. The usefulness of this technique as a measure of vagal activation is discussed.

Stomatal behavior and photosynthetic performance under dynamic light regimes in a seasonally dry tropical rain forest.

Rates of photosynthetic induction upon exposure to high light and rates of induction loss after darkening the leaf were measured in the field for four species of tropical shrubs in the family Rubiaceae. During wet season mornings, stomatal conductance (g s) in the shade prior to induction was generally high enough so that the time course of induction was determined primarily by rates of activation of biochemical processes. During wet season afternoons, however, g s values in the shade tended to be considerably lower and photosynthetic induction following a light increase exhibited a slower time course. In the afternoon, the time course of induction was determined by a combination of stomatal opening time and the rates of activation of light regulated enzymes. Stomatal behavior was also correlated with patterns of induction loss following a transfer from high light to darkness. In the afternoon, maximum g s was lower for all species, and for a given time in the darkness, leaves showed a greater loss of induction in the afternoon than in the morning. During the dry season, maximum g s and average values for g s in the shade were reduced in all species. Along with these shifts in stomatal behavior, reduced rates of photosynthetic induction were observed. In the high-light species, the lower maximum g s values observed during the dry season were also correlated with increased induction loss for a given time in the darkness. For all species, stomatal behavior was affected by season and time of day and, with the exception of wet season mornings, stomata appeared to exert significant control over rates of induction and patterns of induction loss. The results of simulation modeling suggest that the observed seasonal and diurnal changes in rates of induction and induction loss can have significant consequences on sunfleck carbon gain under a dynamic light regime.

Stomatal versus biochemical limitations to dynamic photosynthetic performance in four tropical rainforest shrub species.

Photosynthetic performance under dynamic light regimes was assessed in four different species of tropical shrubs from the family Rubiaceae via field gas exchange measurements conducted on Barro Colorado Island, Panamá. Rates of photosynthetic induction and induction loss were assessed throughout the day in both the wet and dry seasons in order to determine the relative roles of stomata and biochemistry in limiting photosynthetic performance under transient light conditions. A high degree of coordination was observed between stomatal conductance and biochemical capacity for CO2 assimilation during induction. Rates of biochemical and overall photosynthetic induction sharply decreased when initial stomatal conductance fell below a narrow range of critical values. Time of day or season did not affect rates of biochemical deactivation upon shading, but did influence stomatal closure, which often exerted a significant influence over induction loss in the darkness. In measurements of total assimilation due to a 60-s light pulse, both biochemical activity and stomatal conductance were linearly related to total CO2 uptake. Only during the mornings of the wet season was stomatal conductance consistently high enough to be non-limiting to dynamic photosynthetic performance. At all other times, stomatal behavior exercised significant influence over induction times, photosynthetic induction loss, and total CO2 uptake from 60-s light pulses.

Moderators of cardiovascular reactivity to speech: discourse production and group variations in blood pressure and pulse rate.

To explain why speech produces cardiovascular reactivity, an experiment utilized speech tasks varying in self-disclosure and cognitive preparation and measured the blood pressure and pulse rate of African-American and Caucasian, men and women subjects. One hundred and fifty-six college students with a median age of 21 volunteered to participate. The extemporaneous speech task produced lower systolic blood pressure than the prepared speech task. Self-disclosive speech resulted in lower diastolic blood pressure than non-disclosive speech. Females showed higher pulse rate and men exhibited greater diastolic blood pressure while speaking. Results supported previous research suggesting that women are myocardiacal hyperreactors, while men are vascular hypperreactors and suggest that production processes moderate cardiovascular reactivity to speech.

Enhanced cardiovascular and catecholamine responses in women with depressive symptoms.

To test the hypothesis that cardiovascular and sympathetic nervous system responses before and during behavioral stressors are exaggerated among subjects with depressed mood who do not have clinical depressive disorder. Sixty healthy women aged 18-49 were initially asked to complete the Beck Depression Inventory (BDI). The 15 with the highest (Depressive Symptom group) and the 15 with the lowest BDI scores (Control group) underwent stress testing including baseline, postural challenge, a speech task describing responses to a recent anger-arousing experience and recovery. Both systolic and diastolic blood pressure levels were higher in the Depressive Symptom group during baseline, challenges and recovery. This group also showed shorter pre-ejection period (a marker of enhanced cardiac sympathetic activation) and lesser heart rate variability across all test periods. During the speech task only, the Depressive Symptom group exhibited greater increases in plasma norepinephrine and higher cardiac output responses associated with decreased interbeat interval (faster heart rate). These results support the a priori hypothesis regarding enhanced sympathetic and cardiovascular activity. Finally, the BDI scores correlated very highly with lack of perceived emotional social support, reinforcing prior research on the linkage between social isolation and severity of depressive symptoms.

Cardiovascular reactivity to stress and left ventricular mass in youth.

We studied the relationships of cardiovascular reactivity during mental stress with left ventricular mass index in a group of prepubertal children 8 to 10 years old and in a group of peripubertal or postpubertal adolescents 15 to 17 years old. One hundred fifteen participants, varying in age group, sex, and race (black and white), took part in a laboratory stress protocol consisting of a reaction-time task, a mirror tracing task, a cold forehead challenge, and a stress interview. Cardiovascular measures included blood pressure and heart rate, as well as cardiac output, stroke volume, total peripheral resistance, and preejection period obtained noninvasively with impedance cardiography. Measures of left ventricular mass were made by echocardiography. Results indicated that across all participants, left ventricular mass index was associated with cardiovascular responses during the mirror tracing and cold forehead tasks, especially with those responses reflecting increased vasoconstriction. Subgroup analyses showed that these associations were significant for males and sometimes adolescents but not for females and children. As mirror tracing and cold forehead tasks most consistently produce alpha-adrenergic activation, the results suggest a model in which vasoconstriction due to mental stress is related to increased left ventricular mass in susceptible individuals, even at a young age.