Sublethal effects of Texas brown tide on Streblospio benedicti (Polychaeta) larvae

The Texas brown tide bloom is noted for a concordant decline in benthic biomass and species diversity. However, the link between harmful effects induced by Texas brown tide and benthos has not been demonstrated. It has been proposed there may be a larval bottleneck, where larvae, but not adults, suffer adverse effects. This study was performed to test the effect of brown tide alga, Aureoumbra lagunensis, on mortality, growth and behavior of Streblospio benedicti larvae. Growth rates and swimming speeds, but not mortality rates, of polychaete larvae were reduced in cultures with brown tide relative to Isochrysis galbana, which is about the same size as brown tide. Results from this research indicate that brown tide does have harmful sublethal effects for one dominant species of meroplanktonic larvae, which could help explain reduced adult population size.     

Patterns of nuclear genetic variation in the poecilogonous polychaete Streblospio benedicti

The evolution of marine larvae is replete with transitions in trophic mode, but little is known about how, at the genetic level, these transitions are achieved. Basic parameters, including the number of underlying loci, their molecular characteristics, and the population-genetic processes that drive transitions remain unknown. Streblospio benedicti, an abundant benthic polychaete with heritable poecilogony, provides a unique genetically tractable system for addressing these issues. Individuals of S. benedicti vary in diverse aspects of development. Some females produce small, planktotrophic larvae, whereas others produce large, yolky larvae capable of settling without feeding. Here, I present estimates of basic features of nuclear genetic variation in S. benedicti to lay the foundations for subsequent efforts to understand the genetic basis of poecilogony. Sequence of ～20?kb of random nuclear DNA indicates that the nucleotide composition, at 62.1% A?+?T, is typical of lophotrochozoan genomes. Population-genetic data, acquired by sequencing two loci (～2500 bp) in multiple animals of each developmental morph, indicate that the morphs exhibit very little differentiation at random loci. Nucleotide heterozygosity (θπ) is ～0.5-1% per site, and linkage disequilibrium decays within a few kilobases (ρ ～?3?×?10(-3) per site). These data suggest that genetic mapping by association will require a high density of markers, but linkage mapping and identification of regions of elevated inter-morph differentiation hold great promise.     

Development and sexual dimorphism of branchiae in Streblospio benedicti

The Spionidae is one of the largest and most studied annelid families, but to date, the development and differentiation of post-metamorphic anatomy have not been documented. This study used scanning electron microscopy to examine the development of the branchiae, presumed respiratory organs, in Streblospio benedicti. Branchiae in this species are prominent, paired head structures and first appear around the time of metamorphosis, but do not complete their development until the worm reaches the older juvenile or adult stages. We observed that as the branchiae grew, their overall morphology changed through four different shapes: small bud, tubular, tapered, and, finally, bilimbate. In addition, the abfrontal and frontal surfaces each possessed a unique set of cilia patterns, which we named, and these arose in a particular sequence between the 8- and 35-chaetiger stages. This detailed examination of every stage of branchial development led us to discover that branchia in Streblospio benedicti was a sexually dimorphic organ. Streblospio benedicti is one of approximately eight Spionidae in which there is any type of structural sexual dimorphism, and it is the only species in which sexually dimorphic branchiae are found. The male's frontal surface had four unique cilia patterns, and we hypothesize that those located around the medial protrusion capture and control the release of the spermatophores. This first documentation of a spionid's branchial developmental sequence revealed that not only is this respiratory organ involved in reproduction, but it significantly differentiates after metamorphosis through adulthood.     

Fitness Consequences of Maternal Effects in Streblospio benedicti (Annelida: Polychaeta)

SYNOPSIS. The degree to which a female partitions resourcesbetween fecundity and per offspring investment is a centralquestion in life-history theory. Maternal effects may influencethe nature of this tradeoff through their effect on per offspringinvestment and subsequent offspring fitness. The purpose ofthis study was to determine the effect of female age and sizeon brood size (number of offspring), per offspring investment,and fitness in the polychaete Streblospio benedicti. Early stageembryos were collected from brooding females of known age andsize over a period of 100 days; these embryos were counted andanalyzed for their C and N content. Female size had a positiveeffect on brood size; larger females produced larger broods.However, brood size decreased with female age (females did notincrease in size after reaching sexual maturity). Brood sizedeclined 20–46% between 60 and 160 days of age. Duringthis same age period per offspring investment, measured in termsof C and N, increased by 25%. Offspring survivorship and sizeat two weeks post-release from the female were used as measuresof offspring fitness. Offspring survivorship increased 28% between60 and 160 days of age. Increased growth in offspring from olderfemales resulted in a 23% increase in offspring size at twoweeks. Including the maternal age effect in two population modelsfor S. benedicti increased population growth rate (). Populationgrowth was increased to a greater degree when the maternal effectwas modeled by enhancing offspring survival compared to whenfecundity was increased by the same proportional amount. Thissuggests that the maternal effect may be adaptive, particularlywhen conditions for offspring survival and growth are poor.     

Consequences of a poecilogonous life history for genetic structure in coastal populations of the polychaete Streblospio benedicti

In many species, alternative developmental pathways lead to the production of two distinct phenotypes, promoting the evolution of morphological novelty and diversification. Offspring type in marine invertebrates influences transport time by ocean currents, which dictate dispersal potential and gene flow, and thus has sweeping evolutionary effects on the potential for local adaptation and on rates of speciation, extinction and molecular evolution. Here, we use the polychaete Streblospio benedicti to investigate the effects of dimorphic offspring type on gene flow and genetic structure in coastal populations. We use 84 single nucleotide polymorphism (SNP) markers for this species to assay populations on the East and West Coasts of the United States. Using these markers, we found that in their native East Coast distribution, populations of S. benedicti have high‐population genetic structure, but this structure is associated primarily with geographic separation rather than developmental differences. Interestingly, very little genetic differentiation is recovered between individuals of different development types when they occur in the same or nearby populations, further supporting that this is a true case of poecilogony. In addition, we were able to demonstrate that the recently introduced (~100 ya) West Coast populations probably originated from a lecithotrophic population near Delaware.     

Morphogenesis and phenotypic divergence in two developmental morphs of Streblospio benedicti (Annelida,Spionidae)

Abstract. The morphology of marine invertebrate larvae is strongly correlated with egg size and larval feeding mode. Planktotrophic larvae typically have suites of morphological traits that support a planktonic, feeding life style, while lecithotrophic larvae often have larger, yolkier bodies, and in some cases, a reduced expression of larval traits. Poecilogonous species provide interesting cases for the analysis of early morphogenesis, as two morphs of larvae are produced by a single species. We compared morphogenesis in planktotrophic and lecithotrophic morphs of the poecilogonous annelid Streblospio benedicti from the trochophore stage through metamorphosis, using observations of individuals that were observed alive, with scanning electron microscopy, or in serial sections. Offspring of alternate developmental morphs of this species are well known to have divergent morphologies in terms of size, yolk content, and the presence of larval bristles. We found that some phenotypic differences between morphs occur as traits that are present in only one morph (e.g., larval bristles, bacillary cells on the prostomium and pygidium), but that much of the phenotypic divergence is based on heterochronic changes in the differentiation of shared traits (e.g., gut and coelom). Tissue and organ development are compared in both morphs in terms of their structure and ontogenetic change throughout early development and metamorphosis.     

Evolutionary implications of morphogenesis and molecular patterning of the blind gut in the planarian Schmidtea polychroa

The formation of a through-gut was a key innovation in the evolution of metazoans. There is still controversy regarding the origin of the anus and how it may have been either gained or lost during evolution in different bilaterian taxa. Thus, the study of groups with a blind gut is of great importance for understanding the evolution of this organ system. Here, we describe the morphogenesis and molecular patterning of the blind gut in the sexual triclad Schmidtea polychroa. We identify and analyze the expression of goosecoid, commonly associated with the foregut, and the GATA, ParaHox and T-box genes, members of which commonly are associated with gut regionalization. We show that GATA456a is expressed in the blind gut of triclads, while GATA456b is localized in dorsal parenchymal cells. Goosecoid is expressed in the central nervous system, and the unique ParaHox gene identified, Xlox, is detected in association with the nervous system. We have not isolated any brachyury gene in the T-box complement of S. polychroa, which consists of one tbx1/10, three tbx2/3 and one tbx20. Furthermore, the absence of genes like brachyury and caudal is also present in other groups of Platyhelminthes. This study suggests that GATA456, in combination with foxA, is a gut-specific patterning mechanism conserved in the triclad S. polychroa, while the conserved gut-associated expression of foregut, midgut and hindgut markers is absent. Based on these data and the deviations in spiral cleavage found in more basal flatworms, we propose that the lack of an anus is an innovation of Platyhelminthes. This may be associated with loss of gut gene expression or even gene loss.     

Persistent ancestral feeding structures in nonfeeding annelid larvae

Evolutionary loss of the requirement for feeding in larvae of marine invertebrates is often followed by loss of structures involved in capturing and digesting food. Studies of echinoderms suggest that larval form evolves rapidly in response to loss of the requirement for feeding, but a lack of data from other taxa makes it difficult to assess the generality of this result. I show that many members of a large clade of annelids, the Sabellidae, retain ancestral systems for particle capture despite loss of the need and ability to feed. In at least one species, Schizobranchia insignis, an opposed-band system of prototrochal, food-groove, and metatrochal ciliary bands can concentrate suspended particles and transport them to the mouth, but captured particles are invariably rejected because larvae lack a functional gut. The persistence of particle capture systems in larvae of sabellids suggests that they have lost larval feeding very recently, that opposed bands are inexpensive to construct and operate, or that opposed bands have some alternative function. These observations also suggest a hypothesis on how the ability to feed is lost in larvae of annelids and other spiralians following increases in egg size.     

Opposed bands of cilia in the nonfeeding larvae of the serpulid annelid Salmacina tribranchiata

The nonfeeding planktonic larvae of marine invertebrates typically lack larval feeding structures. One puzzling exception to this generalization is the annelid clade Sabellidae, in which nonfeeding larvae possess ciliary bands (specifically, food groove and metatroch) that, to the best of our knowledge, have no function other than in feeding. Nishi and Yamasu (1992b, Bulletin of the College of Sciences, University of the Ryukyus, 54 , 107–121) published a scanning electron micrograph showing that nonfeeding larvae of the serpulid annelid Salmacina dysteri also possess food groove and metatrochal cilia. Here I demonstrate that nonfeeding larvae of Salmacina tribranchiata also bear ciliary bands identifiable as food groove and metatroch by position. High‐speed video of ciliary beat patterns shows that, together with the prototrochal cilia, these bands function in an opposed band system. The presence of feeding structures in nonfeeding annelid larvae is thus more widely distributed than previously recognized. The presence of feeding structures may make evolutionary transitions to planktotrophy more likely, and may underlie an inferred origin of larval feeding in the common ancestor of one of the two major clades of serpulid annelids, Serpulinae.     

Evolutionary relationships of the gut hormones.

Peptides identical or related to mammalian gut hormones occur widely, not just in gut endocrine cells but also in central or peripheral nerves, amphibian skin glands, and a variety of invertebrate tissues. The dual distribution in brain and gut was probably already established early in the vertebrate line; representatives of the oldest vertebrate group, the cyclostomes, have cholecystokinin-like factors in gut endocrine cells and in brain. The related sequences of certain gut peptides, notably gastrin and cholecystokinin (CCK), and secretin, glucagon, vasoactive intestinal polypeptide (VIP), and gastric inhibitory peptide (GIP), indicate evolution from common ancestral molecules by gene duplication and divergence. Functionally important residues are conserved. Thus the COOH-terminal pentapeptide common to gastrin and CCK also contains their minimal active fragment. There are also evolutionary changes at the level of the target organ receptor mechanisms: these are also evolutionary changes at the level of the target organ receptor mechanisms; these are illustrated by evidence suggesting that secretin regulates the flow of pancreatic juice in mammals whereas the structurally related peptide VIP has a similar role in birds.     

Projected timing of perceivable changes in climate extremes for terrestrial and marine ecosystems

Human and natural systems have adapted to and evolved within historical climatic conditions. Anthropogenic climate change has the potential to alter these conditions such that onset of unprecedented climatic extremes will outpace evolutionary and adaptive capabilities. To assess whether and when future climate extremes exceed their historical windows of variability within impact‐relevant socioeconomic, geopolitical, and ecological domains, we investigate the timing of perceivable changes (time of emergence; TOE) for 18 magnitude‐, frequency‐, and severity‐based extreme temperature (10) and precipitation (8) indices using both multimodel and single‐model multirealization ensembles. Under a high‐emission scenario, we find that the signal of frequency‐ and severity‐based temperature extremes is projected to rise above historical noise earliest in midlatitudes, whereas magnitude‐based temperature extremes emerge first in low and high latitudes. Precipitation extremes demonstrate different emergence patterns, with severity‐based indices first emerging over midlatitudes, and magnitude‐ and frequency‐based indices emerging earliest in low and high latitudes. Applied to impact‐relevant domains, simulated TOE patterns suggest (a) unprecedented consecutive dry day occurrence in >50% of 14 terrestrial biomes and 12 marine realms prior to 2100, (b) earlier perceivable changes in climate extremes in countries with lower per capita GDP, and (c) emergence of severe and frequent heat extremes well‐before 2030 for the 590 most populous urban centers. Elucidating extreme‐metric and domain‐type TOE heterogeneities highlights the challenges adaptation planners face in confronting the consequences of elevated twenty‐first century radiative forcing.     

Heterotypic cellular cooperation in gut morphogenesis and differentiation

The development of the gastrointestinal tract, like that of other organs, results from the association of intrinsic genetic endowment, endogenous regulatory mechanisms and environmental influence. The present chapter emphasizes the role of epithelial-mesenchymal interactions during development and crypt-villus migration. Arguments are given as to the role of basement membrane molecules as mediators for these cell interactions. Differential developmental expression of the various extracellular matrix components, which precedes the onset of differentiation markers, suggests that each of these molecules assumes a specific task. Yet, much work still remains to be done before knowing exactly how transfer of morphogenetic signals between epithelial and mesenchymal tissues occurs.     

Phospholipase activity in gut juice of lepidopterous larvae

Phospholipase A activity has been identified in gut juice from larvae of Pieris brassicae, Spodoptera exiguens, and Trichoplusia ni. The specific activity is high and suggests a rôle in lipid breakdown and assimilation.