Larval activity levels and delayed metamorphosis affect post-larval performance in the colonial, ascidian Diplosoma listerianum

It is becoming widely recognized that extending the larval period of marine invertebrates, especially of species with non-feeding larvae, can affect post-larval performance. As these carry-over effects are presumed to be caused by the depletion of larval energy reserves, we predicted that the level of larval activity would also affect post-larval performance. This prediction was tested with the cosmopolitan colonial ascidian Diplosoma listerianum in field experiments in southern Australia. Diplosoma larvae, brooded in the parent colony, are competent to settle immediately after spawning, and they remain competent to metamorphose for > 15 h. Some larvae were induced to metamorphose 0 to 6 h after release, whilst others were induced to swim actively by alternating light and dark periods for up to 3 h prior to metamorphosis. Juvenile colonies were then transplanted to a subtidal field site in Port Phillip Bay and left to grow for up to 3 wk. Extending the larval period and increasing the amount of swimming both produced carry-over effects on post-larval performance. Colonies survived at different rates among experiments, but larval experience did not affect survival rates. Delays in metamorphosis and increased swimming activity did, however, reduce colony growth rates dramatically, resulting in 50% fewer zooids per colony. Moreover, such colonies produced initial zooids with smaller feeding structures, with the width of branchial baskets reduced by 10 to 15%. These differences in branchial basket size persisted and were still apparent in newly budded zooids 3 wk after metamorphosis. Our results suggest that, for D. listerianum, larval maintenance, swimming, and metamorphosis all use energy from a common pool, and increases in the allocation to maintenance or swimming come at the expense of post-larval performance.

The relative energetic costs of the larval period, larval swimming and metamorphosis for the ascidian Diplosoma listerianum

Variation in larval quality has been shown to strongly affect the post-metamorphic performance of a wide range of marine invertebrate species. Extending the larval period of non-feeding larvae strongly affects post-metamorphic survival and growth in a range of species. These 'carry-over' effects are assumed to be due to changes in larval energetic reserves but direct tests are surprisingly rare. Here, we examine the energetic costs ( relative to the costs of metamorphosis) of extending the larval period of the colonial ascidian Diplosoma listerianum. We also manipulated larval activity levels and compared the energy consumption rates of swimming larvae and inactive larvae. Larval swimming was, energetically, very costly relative to either metamorphosis or merely extending the larval period. At least 25% of the larval energetic reserves are available for larval swimming but metamorphosis was relatively inexpensive in this species and larval reserves can be used for post-metamorphic growth. The carry-over effects previously observed in this species appear to be nutritionally mediated and even short (< 3 h) periods of larval swimming can significantly deplete larval energy reserves.

The effect of larval age on morphology and gene expression during ascidian metamorphosis

Metamorphosis is both an ecological and a developmental genetic transition that an organism undergoes as a normal part of ontogeny. Many organisms have the ability to delay metamorphosis when conditions are unsuitable. This strategy carries obvious benefits, but may also result in severe consequences for older larvae that run low on energy. In the marine environment, some lecithotrophic larvae that have prolonged periods in the plankton may begin forming postlarval and juvenile structures that normally do not appear until after settlement and the initiation of metamorphosis. This precocious activation of the postlarval developmental program may reflect an adaptation to increase the survival of older, energy-depleted larvae by allowing them to metamorphose more quickly. In the present study, we investigate morphological and genetic consequences of delay of metamorphosis in larvae of Herdmania momus (a solitary stolidobranch ascidian). We observe significant morphological and genetic changes during prolonged larval life, with older larvae displaying significant changes in RNA levels, precocious migration of mesenchyme cells, and changes in larval shape including shortening of the tail. While these observations suggest that the older H. momus larvae are functionally different from younger larvae and possibly becoming more predisposed to undergo metamorphosis, we did not find any significant differences in gene expression levels between postlarvae arising from larvae that metamorphosed as soon as they were competent and postlarvae developing from larvae that postponed metamorphosis. This recalibration, or convergence, of transcript levels in the early postlarva suggests that changes that occur during prolonged larval life of H. momus are not necessarily associated with early activation of adult organ differentiation. Instead, it suggests that an autonomous developmental program is activated in H. momus upon the induction of metamorphosis regardless of the history of the larva.

Variation in the dispersal potential of non-feeding invertebrate larvae: the desperate larva hypothesis and larval size

For many species of marine invertebrates, variability in larval settlement behaviour appears to be the rule rather than the exception. This variability has the potential to affect larval dispersal, because settlement behaviour will influence the length of time larvae are in the plankton. Despite the ubiquity and importance of this variability, relatively few sources of variation in larval settlement behaviour have been identified. One important factor that can affect larval settlement behaviour is the nutritional state of larvae. Non-feeding larvae often become less discriminating in their 'choice' of settlement substrate, i.e. more desperate to settle, when energetic reserves run low. We tested whether variation in larval size (and presumably in nutritional reserves) also affects the settlement behaviour of 3 species of colonial marine invertebrate larvae, the bryozoans Bugula neritina and Watersipora subtorquata and the ascidian Diplosoma listerianum. For all 3 species, larger larvae delayed settlement for longer in the absence of settlement cues, and settlement of Bugula neritina larvae was accelerated by the presence of settlement cues, independently of larval size. In the field, larger W subtorquata larvae also took longer to settle than smaller larvae and were more discriminating towards settlement surfaces. These differences in settlement time are likely to result in differences in the distance that larvae disperse in the field. We suggest that species that produce non-feeding larvae can affect the dispersal potential of their offspring by manipulating larval size and thus larval desperation.

Variable effects of larval size on post-metamorphic performance in the field

Larval quality may be capable of explaining much of the variation in the recruitment and subsequent population dynamics of benthic marine invertebrates. Whilst the effects of larval nutritional condition on adult performance have received the most attention, recent work has shown that larval size may also be an important and ubiquitous source of variation in larval quality. We examined the effects of variation in larval size on the post-metamorphic survival and growth of Watersipora subtorquata in 2 very different habitats - experimental substrata and pier pilings. We found strong effects of larval size on colony performance, although these varied among experiments. For colonies on experimental substrata, larval size positively affected adult survival and, initially, growth. However, after 3 wk in the field, there was no relationship between larval size and colony size, possibly because colonies were completely surrounded by newly settled organisms. Larval size also positively affected post-metamorphic growth of colonies on pier pilings, but, surprisingly, colonies that came from larger larvae had lower survival than colonies from smaller larvae. Overall, variation in larval size will strongly affect the recruitment and subsequent performance of adults in this species, although this may vary among different habitats. This study highlights the importance of examining the effects of larval quality on adult performance in as realistic conditions as possible, because of the strong interaction between larval size effects and the environment.

Less inhibited with age? Larval age modifies responses to natural settlement inhibitors

As larvae of marine invertebrates age, their response to settlement cues can change. This change can have significant consequences to both the ecology of these organisms, and to their response to antifouling coatings. This study examines how larval age affects the settlement response of larvae to two naturally derived settlement inhibitors, non-polar extracts from the algae Delisea pulchra and Dilophus marginatus, the former of which contains compounds that are in commercial development as antifoulants. Two species of marine invertebrates with non-feeding larvae were investigated: the bryozoans Watersipora subtorquata and Bugula neritina. Larval age strongly affected larval settlement, with older larvae settling at much higher rates than younger larvae. Despite having strong, inhibitory effects on young larvae, the non-polar extracts did not inhibit the settlement of older larvae to the same degree for both species studied. The results show that the effects of ecologically realistic settlement inhibitors are highly dependent on larval age. Given that the age of settling larvae is likely to be variable in the field, such age specific variation in settlement response of larvae may have important consequences for host-epibiont interactions in natural communities.

Complex life cycles and offspring provisioning in marine invertebrates

Offspring size can have pervasive effects throughout an organism's life history. Mothers can make either a few large or many small offspring, and the balance between these extremes is determined by the relationship between offspring size and performance. This relationship in turn is thought to be determined by the offspring's environment. Recently, it has become clear that events in one life-history stage can strongly affect performance in another. Given these strong carryover effects, we asked whether events in the larval phase can change the relationship between offspring size and performance in the adult phase. We manipulated the length of the larval period in the bryozoan Bugula neritina and then examined the relationship between offspring size and various parameters of adult performance under field conditions. We found that despite the adult stage being outplanted into identical conditions, different offspring sizes were predicted to be optimal, depending on the experience of those adults as larvae. This work highlights the fact that the strong phenotypic links between life-history stages may result in optimal offspring size being highly unpredictable for organisms with complex life cycles.

Seawater carbonate chemistry and survival of mollusks: Mercenaria mercenaria, Argopecten irradians, and Crassostrea virginica during experiments, 2009

We present experiments that examined the metamorphosis, growth, and survivorship of larvae from three species of commercially and ecologically valuable shellfish (Mercenaria mercenaria, Argopecten irradians, and Crassostrea virginica) at the levels of CO2 projected to occur during the 21st century and beyond. Under CO2 concentrations estimated to occur later this century (~66 Pa, 650 ppm), M. mercenaria and A. irradians larvae exhibited dramatic declines (>50%) in survivorship as well as significantly delayed metamorphosis and significantly smaller sizes. Although C. virginica larvae also experienced lowered growth and delayed metamorphosis at ~66 Pa CO2, their survival was only diminished at ~152 Pa CO2. The extreme sensitivity of larval stages of shellfish to enhanced levels of CO2 indicates that current and future increases in pelagic CO2 concentrations may deplete or alter the composition of shellfish populations in coastal ecosystems.

Influence of natural settlement cues on the metamorphosis of fiddler crab megalopae, Uca vocator (Decapoda: Ocypodidae)

Megalopas de muitas espécies de caranguejos decápodes aceleram seu período de desenvolvimento até a metamorfose (TOM) quando são expostas a estímulos naturais físicos e/ou químicos característicos do habitat parental. No presente estudo, a influência de estímulos naturais sobre as taxas de muda e sobre o PDM foi investigada nas megalopas do caranguejo violinistac. Os efeitos da (i) lama de diferentes habitats (incluindo habitats de um manguezal bem preservado e de um degradado e poluído) e (ii) 'odores' dos adultos conspecíficos (água do mar acondicionada com caranguejos) sobre a indução da metamorfose foram comparados com (iii) água do mar pura e filtrada (controle). 95 a 100% das megalopas realizaram a metamorfose com sucesso para o primeiro estágio de caranguejo juvenil em todos os tratamentos, incluindo o controle. No entanto, o PDM diferiu significativamente entre os tratamentos. Os estímulos encurtaram significativamente o desenvolvimento, enquanto que a muda foi retardada na ausência deles. O fato de que as megalopas responderam aos estímulos indutores da metamorfose oriundos de ambos os habitats bentônicos dos adultos e de habitats onde eles são ausentes, demonstra que o assentamento nesta espécie pode ocorrer em uma grande variedade de habitats dentro do ecossistema de manguezal, incluindo áreas impactadas.

Neural Mechanisms And Delayed Gastric Emptying Of Liquid Induced Through Acute Myocardial Infarction In Rats.

Background: In pathological situations, such as acute myocardial infarction, disorders of motility of the proximal gut can trigger symptoms like nausea and vomiting. Acute myocardial infarction delays gastric emptying (GE) of liquid in rats. Objective: Investigate the involvement of the vagus nerve, α 1-adrenoceptors, central nervous system GABAB receptors and also participation of paraventricular nucleus (PVN) of the hypothalamus in GE and gastric compliance (GC) in infarcted rats. Methods: Wistar rats, N = 8-15 in each group, were divided as INF group and sham (SH) group and subdivided. The infarction was performed through ligation of the left anterior descending coronary artery. GC was estimated with pressure-volume curves. Vagotomy was performed by sectioning the dorsal and ventral branches. To verify the action of GABAB receptors, baclofen was injected via icv (intracerebroventricular). Intravenous prazosin was used to produce chemical sympathectomy. The lesion in the PVN of the hypothalamus was performed using a 1mA/10s electrical current and GE was determined by measuring the percentage of gastric retention (% GR) of a saline meal. Results: No significant differences were observed regarding GC between groups; vagotomy significantly reduced % GR in INF group; icv treatment with baclofen significantly reduced %GR. GABAB receptors were not conclusively involved in delaying GE; intravenous treatment with prazosin significantly reduced GR% in INF group. PVN lesion abolished the effect of myocardial infarction on GE. Conclusion: Gastric emptying of liquids induced through acute myocardial infarction in rats showed the involvement of the vagus nerve, alpha1- adrenergic receptors and PVN.Fundamento: Distúrbios da motilidade do intestino proximal no infarto agudo do miocárdio podem desencadear sintomas digestivos como náuseas e vômitos. O infarto do miocárdio ocasiona retardo do esvaziamento gástrico (EG) de líquido em ratos. Objetivo: Investigar se existe a influência do nervo vago (VGX), adrenoreceptores α-1, receptores GABAB do sistema nervoso central e participação do núcleo paraventricular (NPV) do hipotálamo no esvaziamento gástrico (EG) e complacência gástrica (CG) em ratos infartados. Métodos: Ratos Wistar (n = 8-15) foram divididos em: grupo infarto (INF), sham (SH) e subdivididos. O infarto foi realizado por ligadura da artéria coronária descendente anterior. A complacência gástrica foi estimada com curvas pressão-volume. Realizada vagotomia por secção dos ramos dorsal e ventral. Para verificar a ação dos receptores GABAB foi injetado baclofeno por via intra ventrículo-cerebral. Simpatectomia química foi realizada com prazosina intravenosa (iv), e na lesão do núcleo paraventricular do hipotálamo foi utilizada corrente elétrica de 1mA/10s, com esvaziamento gástrico determinado por medição da retenção gástrica (% RG) de uma refeição salina. Resultados: Não houve diferença significativa na CG. A vagotomia (VGX) reduziu significativamente a %RG; no grupo INF, o tratamento intra ventrículo-cerebral (ivc) com baclofeno reduziu significativamente a % RG; não houve conclusivamente envolvimento dos receptores GABAB em retardar o EG; o tratamento intravenoso com prazosina reduziu significativamente a %RG no grupo INF. A lesão do NPV aboliu o efeito do infarto do miocárdio no EG. Conclusão: O nervo vago, receptores α-adrenérgicos e núcleo paraventricular estão envolvidos no retardo do esvaziamento gástrico no infarto agudo do miocárdio em ratos.

Delayed postural contraction of transversus abdominis in low back pain associated with movement of the lower limb

The temporal parameters of the response of the trunk muscles associated with movement of the lower limb were investigated in people with and without low back pain (LBP). The weight shift component of the task was completed voluntarily prior to a stimulus to move to allow investigation of the movement component of the response. In the control subjects the onset of electromyographic (EMG) activity of all trunk muscles preceded that of the muscle responsible for limb movement, thus contributing to the feed forward postural response. The EMG onset of transversus abdominis was delayed in the LBP subjects with movement in each direction, while the EMG onsets of rectus abdominis, erector spinae, and oblique abdominal muscles were delayed with specific movement directions. This result provides evidence of a change in the postural control of the trunk in people with LBP.

Extended delayed recall of AVLT word lists: Effects of age and sex on adult performance

An extension of a previous study of age and sex effects on verbal recall (Geffen, Moar, O'Hanlon, lark, & Geffen, 1990) examined forgetting of words over extended delays. The AVLT was administered to 201 normal adults (99 males, 102 females) ranging in age between 20 and 59 years. Recall was tested at intervals of 30 minutes, 24 hours, and 7 days after acquisition. Testing of the latter two intervals was conducted by telephone (Experiment 1, N = 177). After 30 minutes there was no significant loss of the 10 to 11 words retained from five acquisition trials. However, an overall mean of about one word was forgotten after 1 day and a further word after 7 days. The oldest age group (50-59 years) acquired fewer words and forgot more words than the younger groups. Females of all age groups performed slightly better than males at acquisition, at retention, and at recall after longer delays. A second experiment showed that telephone testing at the longer delay intervals was equivalent to testing face to face. These results extend the use of the AVLT by assessing memory decay beyond the immediate testing period. Telephone follow-up is a convenient and economical method of testing delayed recall.

Experimental control of single-mode laser chaos by using continuous, time-delayed feedback

Control of chaos in the single-mode optically pumped far-infrared (NH3)-N-15 laser is experimentally demonstrated using continuous time-delay control. Both the Lorenz spiral chaos and the detuned period-doubling chaos exhibited by the laser have been controlled. While the laser is in the Lorenz spiral chaos regime the chaos has been controlled both such that the laser output is cw, with corrections of only a fraction of a percent necessary to keep it there, and to period one. The laser has also been controlled while in the period-doubling chaos regime, to both the period-one and -two states.

Inhibition of settlement and metamorphosis of the ascidian Herdmania curvata by non-geniculate coralline algae

The surfaces of non-geniculate coralline algae (NCA) are known to induce the settlement and metamorphosis of disparate marine taxa. In this study we investigate the responsiveness of larvae of Herdmania curvata (Ascidiacea: Stolidobranchia) to three species of NCA (Neo-goniolithon brassica-florida, Hydrolithon onkodes, and Lithothamnium prolifer) that cohabit the slope and crest of Heron Reef, Great Barrier Reef. H. curvata larvae were first exposed to these NCA at or within 2 h of hatching, which is 1 to 2 h prior to attaining competence, and then cultured continuously with the NCA for 12 to 14 h. Rates of settlement and metamorphosis of H, curvata cultured in laboratory chambers in the presence of the different NCA were significantly lower than spontaneous rates in seawater. The limited settlement in treatments containing NCA were confined entirely to the chamber periphery, and settlement never occurred on the surface of the NCA. The inhibitory effect was dose-dependent and was stronger in H. brassica-florida and H. onkodes than in L. prolifer. Larvae that did not settle in treatments with NCA had rounded anterior trunks and, in extreme cases, kinked tails with rounded and dissociated tail muscle cells. In some individuals, we observed the anterior chemosensory papillae being sloughed off the larval body. Morphological analysis of trunk ectodermal and mesenchymal nuclei of larvae cultured in the presence of the NCA revealed that general necrotic cell death was occurring. Importantly, H. curvata larvae that were exposed to NCA could not subsequently be induced to metamorphose in KCl-elevated seawater, whereas larvae not exposed to NCA metamorphosed at high rates in KCl-elevated seawater.