Sequencing of behaviors in Aplysia fasciata: integration of feeding, reproduction, and locomotion.

To begin studying the neural basis of higher-order decision-making in Aplysia fasciata, we examined individual bouts of behavior in various conditions of access to food and/or mates. We then determined rules governing transitions between bouts. The data indicate that a single intermediate condition, moving in place, may be centrally related to transitions between behaviors. In all conditions, over 85% of all transitions between behaviors were via moving in place. Moving in place tended to precede and follow other categories of activity (crawling, swimming, immobile), and feeding. Also, moving in place apparently represents a fixed proportion of all bouts of behavior. In each condition, moving in place represented approximately 40% of all bouts, while the number of bouts of other behaviors varied markedly. After a bout of moving in place there was a strong tendency for the animal to return to the behavior performed before moving in place. Additional preferred sequences of behavior were also seen. Courtship tended to precede mating, and crawling preceded swimming.     

Daily variation of multiple behaviors in Aplysia fasciata: integration of feeding, reproduction, and locomotion

Daily variations in feeding, reproductive behaviors, and activity level were examined under three conditions: (1) animals had access to food and mates; (2) there was access only to food; (3) neither food nor mates were present. Behaviors differed in amplitude of variations from their daily mean value. Egg-laying had the strongest tendency to be clustered. Amplitude of variations in courtship and swimming was larger than those of immobility, crawling, and feeding, while moving in place was the least clustered behavior. Changes in state affected the tendency to be clustered. When food and mates were present, amplitude of variations became progressively larger for immobility, moving in place, crawling, and swimming. Behaviors differed in the degree to which variations were patterned as a 1/day oscillation. Relatively little of the variability in crawling, moving in place, courtship, and egg-laying was due to a 1/day oscillation under any condition. By contrast, a large proportion of the variability of immobility, swimming, mating, and eating was modulated by a 1/day oscillation in at least one condition. The contribution of a 1/day oscillation to variability differed in the conditions examined. For immobility and swimming, when food and mates were present, the 1/day component of variability became smaller. By contrast, presence of mates led to an increase in the 1/day component of variability modulating feeding. Daily peaks of swimming and feeding were in phase with one another, and out of phase with mating and feeding. The ratio of locomotion to inactivity is constant, independent of change in external conditions. However, this ratio varied throughout the day. When food and mates were absent, variations of immobility and swimming were strongly related to variations in the inactivity ratio. When food and mates were added, this tendency decreased. The data suggest that daily variations in occurrence of behaviors can be accounted for by interactions between oscillators affecting Aplysia behavior and motivational variables. A 1/day oscillator has strong effects on mating, swimming, and immobility. The effect of the oscillator is modified by changes in motivational state.     

Effects of conspecifics on feeding in Aplysia fasciata

The presence of other Aplysia in the environment increases the time spent feeding. This finding is consistent with the hypothesis that a common arousal mechanism modulates feeding and reproductive behaviors. The arousal level is presumably increased by pheromones released by additional animals.     

Effects of food deprivation upon behavioral patterns and time budgeting of Aplysia fasciata

Effects of food deprivation upon time devoted to different behaviors in Aplysia fasciata were investigated. Time invested in mating, egg-laying, swimming, and movement in place was increased, and time spent immobile and crawling was decreased. Time of occurrence of behaviors was highly synchronized, but differed from that seen when food was available. Changes in behavior seem to be adaptive to conditions in which food is scarce: increased motility may lead to finding sites where food is present, while increased egg-laying and mating may ensure survival of progeny. Behavioral changes can be readily accounted for by removal of postingestive stimuli signaling satiation.     

Activity patterns and time budgeting of Aplysia fasciata under field and laboratory conditions

Activity patterns of Aplysia fasciata were observed in a protected port environment and in an aquarium. In both, major activities were feeding and mating, which collectively took up about 45% of the total time of the animals. Active behaviors occurred primarily at night; much of the day was spent in the inactive state. Activities were highly synchronized, with large numbers of animals performing the same behaviors simultaneously at a specific time. Mating and eating occurred primarily at different times; relatively few animals were observed performing these behaviors simultaneously. Many animals laying eggs were simultaneously mating as females.     

Common regulation of feeding and mating in Aplysia fasciata: pheromones released by mating and by egg cordons increase feeding behavior.

We examined whether pheromones released by reproductive behaviors (mating and egg-laying) affect feeding behavior. A preliminary experiment demonstrated that the quantity of food eaten can be used to measure the effects of pheromones on feeding. Using this measure, we then showed that Aplysia that were prevented from mating, but that were in the same aquarium as mating conspecifics, eat more food than do Aplysia in a medium lacking mating animals. Mating and feeding were not temporally correlated, indicating that pheromones released by mating probably do not initiate feeding, but rather modulate feeding after it has begun. Aplysia that were in the same aquarium as freshly deposited egg cordons also ate more than did animals in a medium lacking eggs.     

Motivational control of sexual behavior in Aplysia fasciata: sequencing and modulation by sexual deprivation and by addition of partners

Variables similar to those affecting feeding behavior also modulate sexual behavior in Aplysia fasciata, indicating that mating is under motivational control. Motivated behaviors are often patterned into appetitive and consummatory components. Courtship, the appetitive phase of male sexual behavior, was temporally related to subsequent mating. Although many bouts of courtship failed to lead to mating, most incidents of mating were preceded by courtship. Motivation is also characterized by satiation after the goal of the behavior is achieved. We found an increase in likelihood to mate following a period of sexual isolation. Motivated behaviors are also modulated by environmental stimuli that induce arousal. Time spent mating was found to be a function of the number of potential mates accessible for mating. Number of A. fascinata participating in a mating group was also found to be a function of number of potential mates available.     

Effects of partial symmetry, exposure time, and backward masking on judged goodness and reproduction of visual patterns

Six experiments investigated the effects of partial symmetry in visual patterns on judgement of pattern goodness, immediate memory, and learning. In Experiments 1—III pattern goodness ratings were substantially inter-correlated among a self-paced test condition and two conditions producing moderate to severe stimulus degradation (brief tachistoscopic exposure, and backward masking): the less the partial symmetry in a pattern, the lower was the judged goodness of the pattern. In Experiments IV and V immediate reproduction of patterns was observed respectively following exposures of 5-5000 ms, and backward masking. Correct reproduction improved with degree of partial symmetry. Concordant results were found in a free recall learning task (Experiment VI). Correlations between goodness and learning and memory performance for discrete patterns were always substantial. The results strongly suggest that pattern goodness can be appraised reliably and accurately with information processing times too short to permit pattern encoding in short-term visual memory. Evaluation of pattern goodness must therefore rest upon early (precategorical) processing of symmetry features.     

Determinants of pigeons' waiting time: Effects of interreinforcement interval and food delay

Four pigeons performed on three types of schedules at short (i.e., 10, 30, or 60 s) interreinforcement intervals: (a) a delay-dependent schedule where interreinforcement interval was held constant (i.e., increases in waiting time decreased food delay), (b) an interreinforcement-interval-dependent schedule where food delay was held constant (i.e., increases in waiting time increased interreinforcement interval), and (c) a both-dependent schedule where increases in waiting time produced increases in interreinforcement interval but decreases in food delay. Waiting times were typically longer under the delay-dependent schedules than under the interreinforcement-interval-dependent schedules. Those under both-dependent schedules for 1 subject were intermediate between those under the other two schedule types, whereas for the other subjects waiting times under the both-dependent procedure were similar either to those under the delay-dependent schedule or to those under the interreinforcement-interval-dependent schedule, depending both on the subject and the interreinforcement interval. These results indicate that neither the interreinforcement interval nor food delay is the primary variable controlling waiting time, but rather that the two interact in a complex manner to determine waiting times.     

The interactions of courtship, feeding, and locomotion in the behavioral hierarchy of the snail Helix aspersa

Starvation both increased the number of snails that exhibited feeding (i.e., increased feeding proclivity) and decreased the latencies of response to a food stimulus (i.e., increased food arousal). Feeding inhibited locomotion, although starvation itself had no effect. Sexual arousal (the intensity of courtship) increased locomotion, but only in the absence of a mating partner. Sexual proclivity (the likelihood of a snail to court with conspecifics) had no effect on locomotion. In general, proclivity and arousal had different effects on behavior. Food deprivation did not alter the preference of sexually aroused snails for sexual stimuli over food stimuli Both starved and fed courting pairs responded to a food stimulus only during periods of low sexual arousal, although when not sexually aroused, starved snails usually increased the amount of time they spent in contact with a food stimulus. These results suggest that courtship can suppress feeding and that the expression of feeding behavior is dependent upon the occurrence of low levels of sexual arousal (time sharing).     

Rapid and persistent suppression of feeding behavior induced by sensitization training in Aplysia

In Aplysia, noxious stimuli induce sensitization of defensive responses. However, it remains largely unknown whether such stimuli also alter nondefensive behaviors. In this study, we examined the effects of noxious stimuli on feeding. Strong electric shocks, capable of inducing sensitization, also led to the suppression of feeding. The use of multiple training protocols revealed that the time course of the suppression of feeding was analogous to that of sensitization. In addition, the suppression of feeding was present only at the time points in which sensitization was expressed. These results suggest that, in Aplysia, noxious stimuli may produce concurrent changes in neural circuits controlling both defensive and nondefensive behaviors.     

Processing of information about location during locomotion: Effects of a concurrent task and locomotion patterns

How maintenance of orientation during locomotion in unfamiliar environments is accomplished was investigated by having subjects walk behind a moving light line (1.12 m/sec) in a dark room and, from a stopping point, numerically estimate direction and distance to the starting point. Two linear distances (5.0–8.4 m) and the angle of direction change (45, 90, and 135 deg) were orthogonally varied as 12 locomotion patterns. Eight high-school students were assigned to each of three conditions, two in which the starting point was invisible, one in which it was visible. In one of the former conditions the subjects counted backwards rapidly during walking. Lower accuracy and longer latency times were found in the counting condition whilst the other two conditions differed only slightly. The results suggested that accurate maintenance of orientation is achieved by recurrent encoding, coordination and decoding of information about direction change and locomotion distance, processes which demand central processing capacity and therefore are interfered with by a concurrent task.     

Temporal integration of monocular images separated in time: stereopsis, stereoacuity, and binocular luster

We evaluated stereopsis and binocular luster using electronically controlled shutter glasses with alternating monocular stimulation. In Experiment 1, we used the standard method for testing stereoacuity to obtain a gradual measure of stereopsis. Stereo thresholds decreased with increasing alternating frequency of two monocular half-images without a delay between them. Increasing delays led to increasing thresholds. In Experiment 2, we compared stereopsis resulting from two monocular half-images of a random-dot stereogram and binocular luster with respect to the minimum alternating frequency of the two half-images and the maximum interocular delay that were tolerated without a breakdown of the impression. Below 3 Hz, no stereopsis occurred. Binocular luster was observed only above 10 Hz. The mean threshold of interocular delay for detecting the global figure in a random-dot stereogram was about 51 msec, but for binocular luster it was about 20 msec. Overall, temporal integration was better for stereopsis than for binocular luster.     

Processing time of contour integration: the role of colour, contrast, and curvature

We investigated the temporal properties of the red - green, blue-yellow, and luminance mechanisms in a contour-integration task which required the linking of orientation across space to detect a 'path'. Reaction times were obtained for simple detection of the stimulus regardless of the presence of a path, and for path detection measured by a yes/no procedure with path and no-path stimuli randomly presented. Additional processing times for contour integration were calculated as the difference between reaction times for simple stimulus detection and path detection, and were measured as a function of stimulus contrast for straight and curved paths. We found that processing time shows effects not apparent in choice reaction-time measurements. (i) Processing time for curved paths is longer than for straight paths. (ii) For straight paths, the achromatic mechanism is faster than the two chromatic ones, with no difference between the red-green and blue-yellow mechanisms. For curved paths there is no difference in processing time between mechanisms. (iii) The extra processing time required to detect curved compared to straight paths is longest for the achromatic mechanism, and similar for the red - green and blue-yellow mechanisms. (iv) Detection of the absence of a path requires at least 50 ms of additional time independently of chromaticity, contrast, and path curvature. The significance of these differences and similarities between postreceptoral mechanisms is discussed.     

The effects of aging on time reproduction in delayed free-recall

The experiments presented here demonstrate that normal aging amplifies differences in time production occurring in delayed free-recall testing. Experiment 1 compared the time production ability of two healthy aged groups as well as college-aged participants. During the test session, which followed a 24-h delay and omitted all feedback and examples of the two target intervals, the two groups of aged participants' over-produced a 6s interval. This effect is similar in form to errors shown by young participants, but twice the magnitude. Productions of a 17 s interval were generally accurate overall. However, further analysis indicated that the majority of aged participants over-produced the 17 s interval while a minority greatly under-produced the 17 s interval. Furthermore, aged participants showed violations of the scalar property of timing variability in the training session, while in the test session, only those who under-produced the 17 s interval showed this tendency. In contrast, training session performance was good for all participants. Experiments 2 and 3 investigated the ability of the participants in Experiment 1 to reproduce the length of a line from memory, under conditions analogous to those of the time production experiments. These experiments provided tests of the specificity of the errors observed in Experiment 1. Performance in the older participants was accurate, if more variable, compared to the young participants, in contrast to the time production results, indicating that production inaccuracy in free-recall is specific to interval timing in the current context.     

Sex similarities and differences in preferences for short-term mates: what, whether, and why

Are there sex differences in criteria for sexual relationships? The answer depends on what question a researcher asks. Data suggest that, whereas the sexes differ in whether they will enter short-term sexual relationships, they are more similar in what they prioritize in partners for such relationships. However, additional data and context of other findings and theory suggest different underlying reasons. In Studies 1 and 2, men and women were given varying "mate budgets" to design short-term mates and were asked whether they would actually mate with constructed partners. Study 3 used a mate-screening paradigm. Whereas women have been found to prioritize status in long-term mates, they instead (like men) prioritize physical attractiveness much like an economic necessity in short-term mates. Both sexes also show evidence of favoring well-rounded long- and short-term mates when given the chance. In Studies 4 and 5, participants report reasons for having casual sex and what they find physically attractive. For women, results generally support a good genes account of short-term mating, as per strategic pluralism theory (S. W. Gangestad & J. A. Simpson, 2000). Discussion addresses broader theoretical implications for mate preference, and the link between method and theory in examining social decision processes.