Evolutionarily stable sexual allocation by both stressed and unstressed potentially simultaneous hermaphrodites within the same population.

Factors influencing allocation of resources to male and female offspring continue to be of great interest to evolutionary biologists. A simultaneous hermaphrodite is capable of functioning in both male and female mode at the same time, and such a life-history strategy is adopted by most flowering plants and by many sessile aquatic animals. In this paper, we focus on hermaphrodites that nourish post-zygotic stages, e.g. flowering plants and internally fertilising invertebrates, and consider how their sex allocation should respond to an environmental stress that reduces prospects of survival but does not affect all individuals equally, rather acting only on a subset of the population. Whereas dissemination of pollen and sperm can begin at sexual maturation, release of seeds and larvae is delayed by embryonic development. We find that the evolutionarily stable strategy for allocation between male and female functions will be critically dependent on the effect of stress on the trade-off between the costs of male and female reproduction, (i.e. of sperm and embryos). Thus, we identify evaluation of this factor as an important challenge to empiricists interested in the effects of stress on sex allocation. When only a small fraction of the population is stressed, we predict that stressed individuals will allocate their resources entirely to male function and unstressed individuals will increase their allocation to female function. Conversely, when the fraction of stress-affected individuals is high, stressed individuals should respond to this stressor by increasing investment in sperm and unstressed individuals should invest solely in embryos. A further prediction of the model is that we would not expect to find populations in the natural world where both stressed and unstressed individuals are both hermaphrodite.

Clonal structure of the introduced freshwater snail Potamopyrgus antipodarum (Prosobranchia: Hydrobiidae), as revealed by DNA fingerprinting.

Multi-locus DNA fingerprints were obtained from individuals of the hydrobiid snail, Potamopyrgus antipodarum (= P. jenkinsi), by using an RNA derivative (pSPT 18.15) of Jeffrey's 33.15 minisatellite core sequence. Whole-body homogenization of snails yielded 3.21 +/- 0.09 micrograms DNA per individual, producing complex profiles comprising 12-22 fragments within the 1.0-20.0 kilobase (kb) size range. Fingerprints from natural and experimental populations identified three distinct clonal genotypes corresponding to morphological strains A, B and C, with only rare mutational variants. Mother-offspring comparisons of genetic fingerprints revealed genetic stability during apomictic parthenogenesis. Data support the notion that British populations of P. antipodarum comprise three widespread obligate parthenogenetic clones resulting from a mid-19th Century introduction from Australasia. The present-day low levels of genotypic diversity are discussed in relation to the typical occurrence of P. antipodarum in man-made or immature habitats.

Water-borne sperm trigger vitellogenic egg growth in two sessile marine invertebrates.

A diverse array of sessile marine invertebrates mate by passive dispersal of sperm which fertilize the brooded eggs of neighbours. In two such species, a sea-mat (phylum Bryozoa) and an ascidian (phylum Chordata), vitellogenic egg growth is absent in reproductively isolated specimens, but is triggered by a water-borne factor released by conspecifics. In both of these colonial, hermaphroditic species, the active factor can be removed from water by filtration. The effect involves self-/non-self-recognition: water conditioned by a separate subcolony of the same genetic individual does not prompt oocyte growth. In each species, allosperm move from the surrounding water to the ovary and are then stored in close association with the growing oocytes. We concluded that sperm themselves are the water-borne factor that triggers the major phase of female reproductive investment. This mechanism is, to our knowledge, previously undescribed in animals, but has parallels with the initiation of maternal investment in flowering plants following the receipt of compatible pollen. The species studied may be representative of many other aquatic invertebrates which mate in a similar way. The stimulation of egg growth by allosperm could lead to intersexual conflict during oogenesis.

Modification of novelty preferences in rats by current and prior treatment with scopolamine and methylscopolamine.

Two experimental paradigms were adopted to compare effects of scopolamine and its quaternary derivative, methylscopolamine, on the behaviour of albino rats in an exploration box comprising novel and familiar halves. Subjects tested with the first paradigm were exposed to one of the halves, injected and then observed 20 min later. Although both drugs reduced preferences for the previously inaccessible novel half, only scopolamine decreased rearing and increased ambulation. With the second paradigm, behaviour was assessed without any current drug influence. On the 2 days prior to testing the rats had been exposed to one half of the apparatus while drugged. Prior treatment with both scopolamine and methylscopolamine reduced novelty preference to the extent that the familiar half of the apparatus was preferred. Both drugs also reduced rearing (for females only) and ambulation. It was concluded that the results with both paradigms provided some support for the view that reductions in novelty preference by anticholinergic drugs arise from their aversive peripheral actions.

Open-field behaviour in sheep following treatment with the neuroleptics azaperone and acetylpromazine.

The behavior of individual Romney ewes in an open field was observed after intramuscular injection of isotonic saline or two doses of azaperone (0.5, 1.0 mg/kg) or acetylpromazine (0.1, 0.25 mg/kg). Azaperone significantly reduced the distance travelled, urination, and the time spent walking. However it increased nosing of the walls and floor (investigation). Azaperone also increased the numbers of sheep seen grazing and pawing and decreased the number heart vocalizing at least once. The only effect acetylpromazine had was to reduce the number of sheep heart vocalizing. The main results were interpreted as supporting a superior reduction of emotional behaviour by azaperone. It was therefore suggested that, as with pigs, azaperone might be useful for treating stress in sheep.

Effects of adrenalin on novelty choices and activity in rats.

Rearing, ambulation and occupancy of the novel half of an exploration box were observed in Wistar albino rats following intraperitoneal injections of saline, 0.03 or 0.06 mg/kg adrenalin. All three responses were decreased by the drug in a dose-related manner. Adrenalin did not seem to impair ability to discriminate between novel and familiar stimuli since the percentage of rearing that occurred in the novel half was highest in rats who received the 0.06 mg/kg dose. It was concluded that the suppression of novelty choices (and probably rearing and ambulation) by adrenalin was mainly due to its aversive peripheral properties. Pretreatment with 20 mg/kg oxprenolol HCl largely prevented the effects observed earlier, thereby implicating -adrenoceptors in adrenalin's behavioral action.

Cholinergic and monoaminergic substrates of startle habituation.

Research is reviewed arising from the proposition that behavioral habituation is mediated by brain mechanisms operated by the neurotransmitter, acetylcholine. Effects of cholinergic drugs on habituation of the startle response in rats fail to support involvement of acetylcholine. Likewise, serotonergic drug effects do not favor the more recent view that startle habituation depends on brain serotonin, nor is there sufficient evidence for an essential role of either dopamine or noradrenaline . Because of persistence of habituation following challenge with a variety of pharmacological agents, the phenomenon probably depends upon a complex interplay between a number of transmitters and behavioral processes. Contrary to earlier belief, no single transmitter should be seen as crucially responsible for startle habituation.

Effects on open-field behavior of diazepam and buspirone alone and in combination with chronic caffeine.

Effects of diazepam (1, 2 mg/kg) and buspirone (1.25, 2.5 mg/kg) on locomotor and rearing activity were observed in rats tested in an open field. Both doses of each drug reduced ambulation. However, for buspirone, this effect was confined to females. Walking and rearing was reduced by the higher dose of diazepam and rearing by both doses of buspirone. In rats that had ingested approximately 26 mg/kg/day of chronic caffeine for seven days prior to and immediately before testing, all effects of diazepam observed earlier failed to achieve significance except for ambulation. However, all earlier buspirone effects (including female-only decreased ambulation) were unaffected by the caffeine treatment. It was concluded that buspirone may be preferable to diazepam as an anxiolytic when in the presence of regular caffeine ingestion.

Sex-and age-dependent effects of prenatal exposure to caffeine on open-field behavior, emergence latency and adrenal weights in rats.

Pregnant rats were provided with drinking water containing 0, 0.23 or 0.3 mg/ml of caffeine throughout gestation. These concentrations gave rise to daily doses of 0, 28 and 36 mg/kg. Open-field behavior and latencies to emerge from a darkened chamber were observed in offspring at regular intervals from 1 to 8 months after birth. The main results revealed increases in open-field locomotor and rearing activity with 28 but not 36 mg/kg/day. The opposite pattern characterized emergence latency. These changes were more typical of male rats particularly when older. Combining the present results with those of an earlier study by the authors strengthened the curvilinear trends observed and led to the conclusion that, low doses of prenatal caffeine increase activity and decrease emotionality. Higher doses may have the opposite effects to the point that the significant differences from control subjects reported earlier can occur. When 8 months old, female but not male rats prenatally exposed to 36 mg/kg/day of caffeine had significantly heavier adrenal glands than controls.

Behavioral effects of prenatal exposure to caffeine in rats.

For the first ten days of gestation, rats received daily intraperitoneal injections of 10-40 mg/kg of caffeine. Open field behavior of their fostered offspring was observed 61, 145 and 188 days after birth. While there were no obvious physical effects of the prenatal experience, at 61 days caffeine exposure led to an increase in the number of times seen walking for males only and increased ambulation (distance travelled) for both sexes. At 145 days occupancy of centre squares of the apparatus and latencies of emergence from a dark box into an illuminated arena were higher for caffeine-exposed males only. When 188 days old, rats exposed to 20 mg/kg of caffeine tended to exhibit less locomotor activity and more grooming behavior while spending more time in corners of the apparatus. Male rats prenatally exposed to 20 mg/kg of caffeine avoided the centre squares of the apparatus. It was concluded that prenatal caffeine had modified the development of mechanisms controlling voluntary motor activity in the youngest rats. However, at older ages, the prenatal effect was probably manifested as increased timidity or emotional reactivity. Males were often affected differently from females by the prenatal treatment.

Effects of diazepam and cyclohexyladenosine on open-field behavior in rats perinatally exposed to caffeine.

Adult rat offspring earlier exposed to maternally ingested caffeine during both gestation and lactation were observed in an open field following acute administration of diazepam or cyclohexyladenosine. While both drugs reduced measures of locomotion and emotional reactivity, caffeine-exposed rats showed evidence of greater sensitivity to cyclohexyladenosine (but not diazepam) as determined by its effects on grooming behavior and tendencies to occupy the center squares of the apparatus. This suggested that adenosine (A1) rather than benzodiazepine receptor activity had been affected by the perinatal caffeine experience which also reduced locomotor activity while increasing center occupancy. The acute effects of diazepam and cyclohexyladenosine also depended largely on the sex of the subjects. Diazepam affected locomotor activity more and both drugs affected defecation less in females than in males. No other interaction involving sex was significant.

Annual production of two Nova Scotian populations of Nucella lapillus (L.).

One population of Nucella lapillus, under conditions of food shortage, decreased in biomass with an annual production of 5.5 kcal per m2. The second population, with plenty of food, gained biomass and the annual production was 16.7 kcal per m2.Individuals grew faster, while attaining sexual maturity and terminating growth at a larger size in the second population. In both populations, energy produced as gametes exceeded the energy produced due to growth. A 2.56 cm female was estimated to produce 46.6 egg capsules (about 1 kcal) a year but the output of males was unknown and assumed to be half that of females. The sex ratio was equal.

Foraging strategies of dogwhelks, Nucella lapillus (L.): interacting effects of age, diet and chemical cues to the threat of predation.

The effects of diet history, hunger and predation risk on short-term behavioral decisions of dogwhelks were tested in a specially designed test apparatus, termed a linear feeding array (LFA). The LFA consists of a sequential series of prey items mounted in a flume with unidirectional current directed towards a test (predatory) animal, and into which potential olfactory cues regarding predation risk are introduced. For dogwhelks the array was constructed vertically to accomodate intertidal foraging movements and is termed a vertical linear array (VLA). The behaviors exhibited by the dogwhelks were interpreted from distribution patterns in the VLA. Recent experimental studies and advances in optimal foraging theory provided the basis for the hypotheses tested in the VLA, which included: foraging and other behaviors are affected by predation, animals will avoid risk in the presence of predation threat, responses to predation threat will be proportional to the number and kinds of predator cues present, and starved animals will take greater risks than fed animals. We also test the proposition that foraging decisions are further modified by age. Three groups of juvenile and adult animals were maintained on diets of barnacles, mussels or no food (starved). The scent of crabs and damaged conspecifics served as olfactory cues to predation risk. Dogwhelks exhibited a range of behaviors in the VLA including: sheltering, searching, feeding, and aerial climbing. Distribution of animals in the tank assumed a relatively stable pattern after 2-3 h. These patterns were interpreted as the consequence of heirarchial decision making including: (i) a decision to become active, leaving the resting place or water refuge adopted during initial placement, followed by (ii) a decision to move vertically upwards or downwards, and (iii) a decision to attack prey when encountered. Analysis of movement patterns revealed that the initial decision, analogous to leaving a crevice as the tide comes in, was influenced in adults by predator cues and in juveniles by both predator cues and diet history. Perceived risk, as crab and damaged-conspecific odors, made individuals more likely to remain inactive, a risk-avoiding strategy for animals already in a refuge. Starved animals were more likely to descend into the tank and attack prey than fed animals. Our results support the hypotheses that higher-order predators affect the foraging decisions of dogwhelks and that juveniles and satiated animals are more sensitive to predation risk than starved ones. Together, these and earlier studies suggest that dogwhelks assess their environment before foraging, and that they are attuned to reducing the risks of mortality.

Effects of physostigmine on novelty-related location preferences.

Novelty-related location preferences and activity in an exploration box were recorded for male and female Wistar albino rats following intraperitoneal injections of 0.04 or 0.08 mg/kg of either physostigmine or neostigmine. Although rearing was reduced by the highest dose of both drugs and ambulation was reduced by the same dose of neostigmine, neither agent affected the significant preferences for novelty that typified all subjects. In a second experiment designed to assess the effects of 0.08 mg/kg of the two drugs administered during rather than after confinement to the familiar half of the apparatus, neostigmine reduced rearing, walking, and ambulation while increasing defecation, but physostigmine did not affect any response. While some minor motor impairment may have arisen from its peripheral effects, the lack of changes in novelty-related location preferences failed to support facilitation of either novelty avoidance or habituation by physostigmine suggested in previous studies.

Chronic imipramine effects on exploratory behavior in rats.

Approximately 100 days old hooded rats, socially isolated or group-housed since weaning, received 15 daily IP injections of isotonic saline, 10 mg/kg or 20 mg/kg of imipramine HCl. Following their last injection, the rats' active choices of a novel environment, frequencies of rearing and grooming, and cells entered in an exploration box were recorded. The drug treatment reduced rearing, ambulation and (for isolated rats only) grooming, but had no effect on novelty choices. There was a significant weight loss with the higher dose and (for males only) with social isolation during the drug treatment period. While imipramine reduced grooming in isolated but not group-housed rats, there were no other interactions between the two forms of treatment. It was concluded that, in spite of its sedative action on motor activity, chronic imipramine did not alter curiosity about a novel environment.

Scopolamine induced changes in activity and reactions to novelty.

The behavior of hooded rats was observed in an exploration box comprising novel and familiar halves following IP injections of 0.1, 0.25, 0.5, 0.75 or 1.00 mg/kg scopolamine or isotonic saline. Drug administration occurred after, rather than before, exposure to one of the alternative halves. All doses decreased reactions to the previously inaccessible novel half but decreases were greater for the 2 lowest doses. Rearing behavior was also suppressed by each dose whereas the number of apparatus cells entered (locomotion) was decreased by low doses but increased by high. The 3 behavioral measures showed declines in frequency during the course of each experimental session. However, low doses of the drug enhanced and high doses retarded these declines for rearing and cells entered. The study illustrated the difficulty in explaining data by unitary processes (such as attenuated habituation) when several behavioral indices and drug doses are employed within a single investigation.

Aversive stimulus properties of scopolamine.

The drug state produced in rats by intraperitoneal injections of scopolamine hydrobromide (1.2 mg/kg) was treated as a putative aversive US. This US was paired with a distinctive spatial location in a shuttle box for 6 of 12 daily sessions by confining the subject to one side following scopolamine and to the other side following saline (6 sessions). Two groups of 8 subjects each received zero and 20 min post-injection delays respectively. Following zero delay, but not 20 min delay, subjects avoided the side associated with scopolamine in drug-free, free choice tests. This is evidence that the immediate post-injection drug state induced by scopolamine is aversive.

Methylscopolamine and conditioned location avoidance.

On alternate days, rats were confined to one side of a shuttlebox following IP administration of saline and to the other following the peripherally-acting muscarinic antagonist, methylscopolamine (1.2 mg/kg). They later avoided the side associated with the drug effect. By duplicating an earlier finding with centrally- and peripherally-acting scopolamine, this result identified aversive peripheral actions of the two drugs as mainly responsible for the effects observed.

Behavioral effects of exposure to caffeine during gestation, lactation or both.

Open-field behavior and latencies of emergence from a darkened chamber to a brightly lit arena were recorded at 1, 2, 4 and 6 months after birth in male and female rats that had been exposed to 26 or 45 mg/kg/day caffeine ingested by dams in their drinking water during gestation, 25 or 35 mg/kg/day during lactation or to the two low or high doses ingested during both gestation and lactation. One or both of the gestational or lactational doses reduced locomotor activity and increased defecation in the open field at all ages for males only. Rearing was decreased for both sexes by 25 mg/kg/day lactational caffeine. Numbers of rats that failed to or took longer than 1 min to emerge into the brightly lit arena were increased by 26 mg/kg/day gestational caffeine. All rats that had been exposed to either dose combination of caffeine during both gestation and lactation showed less locomotor and rearing activity, reduced tendencies to emerge within 1 min and, at 6 months of age only, more defecation in the open field. It was concluded that the effects of gestational and lactational exposure to caffeine were additive in their modification of the developing brain as reflected in decreased motor activity possibly arising from heightened emotional reactivity to the testing situation. Hypersensitivity of males to caffeine exposure during either gestation or lactation separately seemed to diminish when exposure was increased for all rats through experience of the drug during both gestation and lactation. Possible involvement of caffeine-induced increases in adenosine receptors in the type of results obtained was discussed.

Mechanisms for turn alternation in four invertebrate species.

Free-choice behavior following one or more forced turns was observed in representatives of four invertebrate classes (earthworms, woodlice, millipedes, earwigs). While all animals alternated, species differences occurred in free turn angle and the effects of varied angle and number of forced turns. Overall, woodlice and millipedes turned at sharper angles and responded more to the forced turn conditions than earthworms and earwigs. From behavior observed following three forced turns in one direction and then one in the opposite, it was concluded that, in earlier experiments, earthworms alternated via tactile cues, woodlice mainly used kinesthetic but could also use tactile cues, millipedes mainly used tactile but could also use kinesthetic cues and earwigs may have relied on kinesthetic cues alone. Since phyletic differences did not seem appropriate, the results were discussed in terms of other characteristics such as body shape and life style.