Sylvia Markovich

Behavioral patterns of Aplysia fasciata along the Mediterranean Coast of Israel

Aplysia fasciata were observed in a number of environments. They were usually aggregated. Animals were occasionally buried in sand. Aplysia were less mobile and more deeply hidden when waves were strong. Aplysia swam only in a calm environment. A wide variety of seaweeds were eaten, but the most common food was Ulva lactuca. Food arousal and satiation occur in the field. Often the pattern of feeding was a gradual slowing down and eventual cessation of eating; however, many meals deviated from this pattern. Appetitive behaviors preceding mating as a male were similar to those preceding eating. Aroused animals mated as males, while passive animals mated as females; mating as a male produced arousal, expressed as an increased likelihood to respond to food or to mate as a male. Two males could simultaneously impregnate one female. Animals mate in large groups, constantly changing partners. Mating groups occur in linear chain, branched chain, and closed chain configurations. Egg laying and egg masses were observed in areas inhabited by animals. Inking was never observed without exprimenter intervention, even when a crab attacked an Aplysia.

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.

Parametric features of inhibition of feeding in Aplysia by associative learning, satiation, and sustained lip stimulation

In order to determine whether different classes of behavioral plasticity affect common or unique neural loci, the effects of three types of processes that inhibit feeding in Aplysia were quantified. Changes in feeding behavior due to an associative learning task in which animals learn that food is inedible were compared with behavioral effects caused by satiation and by sustained lip stimulation. The data indicate that each process modifying feeding can be characterized by differences in time to stop responding to food, by differences in specificity of the decrement to a particular food, and by different patterns of motor output before complete cessation of responsiveness. The data suggest each process inhibiting feeding acts at a different neural site. Learning that food is inedible may be due to facilitation of a specific sensory pathway onto pattern generators producing rejection responses. Sustained lip stimulation seems to inhibit feeding by causing a decrement in all outputs of a particular sensory pathway. Finally, satiation appears to represent inhibition of feeding motor elements.

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 food and mates on time budget in Aplysia fasciata: Integration of feeding, reproduction, and locomotion

This study examines the time budgeted to different behaviors in Aplysia fasciata under three conditions: (1) animals have constant access to food and mates: (2) there is access to food, but not to mates; (3) neither food nor mates are present. The data suggest a number of rules underlying behavioral integration: (1) Feeding, reproductive behaviors, and activity seem to be natural categories for behavioral choice. Feeding and reproductive behaviors are controlled in tandem by a common arousal mechanism, while time left over after animals feed and reproduce is distributed in a fixed ratio between locomotion (crawling and swimming) and inactivity (immobility and movement in place). (2) Relative distribution between different forms of locomotion and inactivity is modified by changes in motivational state. More time is spent swimming than crawling when feeding and/or mating is prevented, while more time is spent moving in place than immobile when the arousal level is increased. (3) Feedback control of feeding and reproduction is asymmetric. Satiation of feeding inhibits the common arousal. In the absence of food, time spent on reproductive behaviors increases due to disinhibition of the common arousal. By contrast, positive feedback arising from sexual behavior excites the common arousal. When mating is prevented by removing potential mates, time spent feeding decreases. (4) Generally, animals choose between performing the three main categories of behavior. Although Aplysia simultaneously can feed and mate, or locomote and mate, they do so infrequently. By contrast, different types of reproductive behaviors (male mating, female mating, egg-laying) are commonly performed simultaneously.

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.

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.

ASYMMETRY OF MALE AND FEMALE MATING IN MIXED-SPECIES GROUPS OF APLYSIA FASCIATA POIRET AND APLYSIA BRASILIANA RANG (MOLLUSCA: GASTROPODA)

ABSTRACT Aplysia fasciata and A. brasiliana are found on opposite sides of the Atlantic Ocean, but are morphologically and behaviorally very similar. To determine the relationship between these two species, mating patterns were examined in single-species and mixed-species groups of 2 and 4 individuals. In single—species groups, A. brasiliana spent less time mating than did A. fasciata. However, the relative distribution of time spent mating as a male, as a female, or in both sexual roles simultaneously was virtually identical in both species. The overall time spent mating in mixed-species groups was similar to that seen in single-species groups of A. brasiliana, and was significantly less than in single-species groups of A. fasciata. In mixed-species groups there were relative increases in the likelihood for A. fasciata to mate as males, and for A. brasiliana to mate as females. Our data indicate that when animals with differing sexual drives are in contact, the animal with the stronger sexual drive gener...

CONTROL OF INDIVIDUAL BOUTS OF BEHAVIOR IN APLYSIA FASCIATA: INTEGRATION OF FEEDING, REPRODUCTION, AND LOCOMOTION

ABSTRACT Control of individual bouts of different behaviors was examined in Aplysia fasciata, as part of a long-term project examining the principles underlying higher-order decision-making. Previous studies have shown that access to food and to mates affects the total time devoted to various behaviors. We examined whether these factors also affect either bout lengths or bout frequencies. Access to food inhibited mating, by decreasing both bout lengths and bout frequencies, while access to mates increased feeding by increasing bout length. Bouts of mobility were affected in complex ways by access to food and mates: food had opposite effects on bout lengths and bout frequencies, while mates decreased bout lengths.

Modulation of respiratory pump rate by reproductive behaviors in freely behaving pairs of Aplysia fasciata.

Respiratory pumping in Aplysia is a spontaneously occurring behavior whose neural circuitry has been explored, but whose natural functions are incompletely understood. Respiratory pump rate was examined in freely behaving pairs of Aplysia fasciata, to determine whether it is modified by the occurrence of mating and other behaviors. The background rate of respiratory pumping was approximately 2/hour. This rate was maintained while animals were immobile, moving in place, crawling, or feeding. The rate was increased to over 8/hour during courtship and to approximately 4/hour during female mating and was reduced to approximately 1/hour during male-mating. These data suggest that respiratory pumping has a reproductive function, perhaps in dispersal of pheromones that are released during female-mating and courtship. Respiratory pumping never occurred while animals were swimming, suggesting that respiratory pumping and swimming may be mutually incompatible behaviors. Respiratory pumping was less common by night than by day.