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Extreme low pH events in estuaries and upwelling areas can modulate the phenotypic and genetic diversity of natural populations. To test this hypothesis, we explored the linkage between local scale extreme low pH events, genetic diversity, and variation in fecundity-related traits (body size, egg size, and egg production rate) in the broad-dispersal copepod Acartia tonsa. We assessed genetic and phenotypic characteristics of populations by contrasting extreme low pH environments (upwelling and temperate estuary) in the coastal Southeast Pacific, under natural and experimental conditions. These populations showed significant genetic differentiation with higher diversity in mitochondrial and nuclear loci (encoding mtCOI and 18S rRNA) in the estuarine population. Copepods from this population are exposed to more frequent extreme low pH events (< 7.7), and the adult females exhibit consistent phenotypic variation in body size, egg size, and egg production rate across different cohorts. Experimental acclimation to extreme low pH conditions revealed no significant differences in fecundity-related traits between A. tonsa populations. Although these results partially support our hypothesis, the experimental findings suggest other drivers might also influence phenotypic differences in the local environments.

Coccolithophores are potentially affected by ongoing ocean acidification, where rising CO2 lowers seawater pH and calcite saturation state (Ωcal). Southern Patagonian fjords and channels provide natural laboratories for studying these issues due to high variability in physical and chemical conditions. We surveyed coccolithophore assemblages in Patagonian fjords during late spring 2015 and early spring 2017. Surface Ωcal exhibited large variations driven mostly by freshwater inputs. High-Ωcal conditions (max. 3.6) occurred in the Archipelago Madre de Dios. Ωcal ranged from 2.0–2.6 in the western Strait of Magellan and 1.5–2.2 in the inner channel and was subsaturating (0.5) in Skyring Sound. Emiliania huxleyi was the only coccolithophore widely distributed in Patagonian fjords (> 96 % of total coccolithophores), only disappearing in the Skyring Sound, a semi-closed mesohaline system. Correspondence analysis associated higher E. huxleyi biomasses with lower diatom biomasses. The highest E. huxleyi abundances in Patagonia were in the lower range of those reported in Norwegian fjords. Predominant morphotypes were distinct from those previously documented in nearby oceans but similar to those of Norwegian fjords. Moderately calcified forms of E. huxleyi A morphotype were uniformly distributed throughout Patagonia fjords. The exceptional R/hyper-calcified coccoliths, associated with low Ωcal values in Chilean and Peruvian coastal upwellings, were a minor component associated with high Ωcal levels in Patagonia. Outlying mean index (OMI) niche analysis suggested that pH and Ωcal conditions explained most variation in the realized niches of E. huxleyi morphotypes. The moderately calcified A morphotype exhibited the widest niche breadth (generalist), while the R/hyper-calcified morphotype exhibited a more restricted realized niche (specialist). Nevertheless, when considering an expanded sampling domain, including nearby southeast Pacific coastal and offshore waters, even the R/hyper-calcified morphotype exhibited a higher niche breadth than other closely phylogenetically related coccolithophore species. The occurrence of E. huxleyi in naturally low pH–Ωcal environments indicates that its ecological response is plastic and capable of adaptation.

Anoxic marine zones (AMZs) constitute pelagic systems distinguished from the oxygen minimum zones (OMZs) by the complete absence of detectable oxygen and the accumulation of nitrite in mid-waters. At the top of the oxygen-depleted layer and below the oxycline, nutrients are abundant; light intensity is very much reduced (

The cosmopolitan phytoplankter Emiliania huxleyi contrasts with its closest relatives that are restricted to narrower latitudinal bands, making it interesting for exploring how alternative outcomes in phytoplankton range distributions arise. Mitochondrial and chloroplast haplogroups within E. huxleyi are shared with their closest relatives: Some E. huxleyi share organelle haplogroups with Gephyrocapsa parvula and G. ericsonii which inhabit lower latitudes, while other E. huxleyi share organelle haplogroups with G. muellerae, which inhabit high latitudes. We investigated whether the phylogeny of E. huxleyi organelles reflects environmental gradients, focusing on the Southeast Pacific where the different haplogroups and species co-occur. There was a high congruence between mitochondrial and chloroplast haplogroups within E. huxleyi. Haplogroup II of E. huxleyi is negatively associated with cooler less saline waters, compared to haplogroup I, both when analyzed globally and across temporal variability at the small special scale of a center of coastal upwelling at 30° S. A new mitochondrial haplogroup Ib detected in coastal Chile was associated with warmer waters. In an experiment focused on inter-species comparisons, laboratory-determined thermal reaction norms were consistent with latitudinal/thermal distributions of species, with G. oceanica exhibiting warm thermal optima and tolerance and G. muellerae exhibiting cooler thermal optima and tolerances. Emiliania huxleyi haplogroups I and II tended to exhibit a wider thermal niche compared to the other Gephyrocapsa, but no differences among haplogroups within E. huxleyi were found. A second experiment, controlling for local adaptation and time in culture, found a significant difference between E. huxleyi haplogroups. The difference between I and II was of the expected sign, but not the difference between I and Ib. The differences were small (≤1°C) compared to differences reported previously within E. huxleyi by local adaptation and even in-culture evolution. Haplogroup Ib showed a narrower thermal niche. The cosmopolitanism of E. huxleyi might result from both wide-spread generalist phenotypes and specialist phenotypes, as well as a capacity for local adaptation. Thermal reaction norm differences can well explain the species distributions but poorly explain distributions among mitochondrial haplogroups within E. huxleyi. Perhaps organelle haplogroup distributions reflect historical rather than selective processes.

Cell division in all eukaryotes depends on function of the spindle, a microtubule-based structure that segregates chromosomes to generate daughter cells in mitosis or haploid gametes in meiosis. Spindle size adapts to changes in cell size and shape, which vary dramatically across species and within a multicellular organism, but the nature of scaling events and their underlying mechanisms are poorly understood. Cell size variations are most pronounced in early animal development, as egg diameters range from tens of microns up to millimeters across animal phyla, and decrease several orders of magnitude during rapid reductive divisions. During early embryogenesis in the model organisms X. laevis and C. elegans, the spindle scales with cell size [1, 2], a phenomenon regulated by molecules that modulate microtubule dynamics [3-6], as well as by limiting cytoplasmic volume [7, 8]. However, it is not known to what extent spindle scaling is conserved across organisms and among different cell types. Here we show that in a range of metazoan phyla, mitotic spindle length decreased with cell size across an ∼30-fold difference in zygote size. Maximum spindle length varied, but linear spindle scaling occurred similarly in all species once embryonic cell diameter reduced to 140 μm. In contrast, we find that the female meiotic spindle does not scale as closely to egg size, adopting a more uniform size across species that most likely reflects its specialized function. Our analysis reveals that spindle morphometrics change abruptly, within one cell cycle, at the transition from meiosis to mitosis in most animals.

Cáhuil Lagoon in central Chile harbors distinct microbial communities in various solar salterns that are arranged as interconnected ponds with increasing salt concentrations. Here, we report the metagenome of the 3.0- to 0.2-µm fraction of the microbial community present in a crystallizer pond with 34% salinity.

Cell membrane intact polar lipids (IPLs) are chemotaxonomic biomarkers whose abundances and distributions in water column environments reflect the living biomass of in situ microbial communities, and can be used to determine the relative contribution of distinct functional and phylogenetic groups to water column carbon stocks. The diversity of IPLs in marine environments is, however, vast, while our knowledge of their biological origins remains limited. Here, we study the distribution of IPLs in size-fractionated suspended organic matter from the oxygen minimum zone (OMZ) of the eastern tropical South Pacific (ETSP) off northern Chile. Canonical correspondence analyses of total IPL abundances and water column physiochemistry demonstrate distinct distributions of microbial sources associated with different geochemical regions in the water column (chlorophyll maximum, upper chemocline, lower chemocline, upper OMZ, core OMZ, and mesopelagic region). Furthermore, the distribution of IPLs in free-living (0.3–2.7 μm) and particle-attached (2.7–53 μm) suspended organic matter differs, suggesting distinct biological sources in each size fraction. While some parallels exist, the diversity and distribution of IPLs in the OMZ system of the ETSP off northern Chile exhibited some unique features compared to other OMZ systems; for instance, we observed a significantly lower contribution of betaine lipids from phytoplanktonic sources, possibly reflecting a physiological response to severe N-limitation in this area. The overall IPL abundance in the two size fractions also indicates a dominance of free-living biomass in the OMZ and mesopelagic regions, suggesting that these areas of the water column could provide additional sources of submicrometer-sized organic carbon to deeper waters. This study improves the utility of IPLs as chemotaxonomic biomarkers by providing insight into the contrasting distributions of microbial biomass from different life modes (free-living and particle-attached). Our results suggest that microbial production in low oxygen environments may be more important to total water column carbon stocks than previously thought.

The interplay of coastal oceanographic processes usually results in partial pressures of CO2 (pCO2) higher than expected from the equilibrium with the atmosphere and even higher than those expected by the end of the century. Although this is a well-known situation, the natural variability of seawater chemistry at the locations from which tested organisms or communities originate is seldom considered in ocean acidification experiments. In this work, we aimed to evaluate the role of the carbonate chemistry dynamics in shaping the response of coastal phytoplankton communities to increased pCO2 levels. The study was conducted at two coastal ecosystems off Chile, the Valdivia River estuary and the coastal upwelling ecosystem in the Arauco Gulf. We characterized the seasonal variability (winter/summer) of the hydrographic conditions, the carbonate system parameters, and the phytoplankton community structure at both sites. The results showed that carbonate chemistry dynamics in the estuary were mainly related to seasonal changes in freshwater discharges, with acidic and corrosive conditions dominating in winter. In the Arauco Gulf, these conditions were observed in summer, mainly associated with the upwelling of cold and high pCO2 (>1,000 μatm) waters. Diatoms dominated the phytoplankton communities at both sites, yet the one in Valdivia was more diverse. Only certain phytoplankton groups in this latter ecosystem showed a significant correlations with the carbonate system parameters. When the impact of elevated pCO2 levels was investigated by pCO2 manipulation experiments, we did not observe any significant effect on the biomass of either of the two communities. Changes in the phytoplankton species composition and abundance during the incubations were related to other factors, such as competition and growth phases. Our findings highlight the importance of the natural variability of coastal ecosystems and the potential for local adaptation in determining responses of coastal phytoplankton communities to increased pCO2 levels.

Marine multicellular organisms inhabiting waters with natural high fluctuations in pH appear more tolerant to acidification than conspecifics occurring in nearby stable waters, suggesting that environments of fluctuating pH hold genetic reservoirs for adaptation of key groups to ocean acidification (OA). The abundant and cosmopolitan calcifying phytoplankton Emiliania huxleyi exhibits a range of morphotypes with varying degrees of coccolith mineralization. We show that E. huxleyi populations in the naturally acidified upwelling waters of the eastern South Pacific, where pH drops below 7.8 as is predicted for the global surface ocean by the year 2100, are dominated by exceptionally over-calcified morphotypes whose distal coccolith shield can be almost solid calcite. Shifts in morphotype composition of E. huxleyi populations correlate with changes in carbonate system parameters. We tested if these correlations indicate that the hyper-calcified morphotype is adapted to OA. In experimental exposures to present-day vs. future pCO2 (400 vs. 1200 µatm), the over-calcified morphotypes showed the same growth inhibition (−29.1±6.3 %) as moderately calcified morphotypes isolated from non-acidified water (−30.7±8.8 %). Under the high-CO2–low-pH condition, production rates of particulate organic carbon (POC) increased, while production rates of particulate inorganic carbon (PIC) were maintained or decreased slightly (but not significantly), leading to lowered PIC ∕ POC ratios in all strains. There were no consistent correlations of response intensity with strain origin. The high-CO2–low-pH condition affected coccolith morphology equally or more strongly in over-calcified strains compared to moderately calcified strains. High-CO2–low-pH conditions appear not to directly select for exceptionally over-calcified morphotypes over other morphotypes, but perhaps indirectly by ecologically correlated factors. More generally, these results suggest that oceanic planktonic microorganisms, despite their rapid turnover and large population sizes, do not necessarily exhibit adaptations to naturally high-CO2 upwellings, and this ubiquitous coccolithophore may be near the limit of its capacity to adapt to ongoing ocean acidification.

Abstract Background Nemertean embryos undergo equal spiral cleavage, and prior fate-mapping studies showed that some also exhibit key aspects of spiralian lineage-based fate specification, including specification of the primary trochoblasts, which differentiate early as the core of the prototroch of the spiralian trochophore larva. Yet it remains unclear how the nemertean pilidium larva, a long-lived planktotroph that grows substantially as it builds a juvenile body from isolated rudiments, develops within the constraints of spiral cleavage. Results We marked single cells in embryos of the pilidiophoran Maculaura alaskensis to show that primary, secondary, and accessory trochoblasts, cells that would make the prototroch in conventional spiralian trochophores (1q2, 1q12, and some descendants of 2q), fully account for the pilidium’s primary ciliary band, but without undergoing early cleavage arrest. Instead, the primary ciliary band consists of many small, albeit terminally differentiated, cells. The trochoblasts also give rise to niches of indefinitely proliferative cells (“axils”) that sustain continuous growth of the larval body, including new ciliated band. Several of the imaginal rudiments that form the juvenile body arise from the axils: in particular, we show that cephalic imaginal disks originate from 1a2 and 1b12 and that trunk imaginal disks likely originate from 2d. Conclusions The pilidium exhibits a familiar relation between identified blastomeres and the primary ciliated band, but the manner in which these cells form this organ differs fundamentally from the way equivalent cells construct the trochophore’s prototroch. Also, the establishment, by some progeny of the putative trochoblasts, of indeterminate stem cell populations that give rise to juvenile rudiments, as opposed to an early cleavage arrest, implies a radical alteration in their developmental program. This transition may have been essential to the evolution of a maximally indirect developing larval form—the pilidium—among nemerteans.

Our current understanding of biology is heavily based on a small number of genetically tractable model organisms. Most eukaryotic phyla lack such experimental models, and this limits our ability to explore the molecular mechanisms that ultimately define their biology, ecology, and diversity. In particular, marine protists suffer from a paucity of model organisms despite playing critical roles in global nutrient cycles, food webs, and climate. To address this deficit, an initiative was launched in 2015 to foster the development of ecologically and taxonomically diverse marine protist genetic models. The development of new models faces many barriers, some technical and others institutional, and this often discourages the risky, long-term effort that may be required. To lower these barriers and tackle the complexity of this effort, a highly collaborative community-based approach was taken. Herein, we describe this approach, the advances achieved, and the lessons learned by participants in this novel community-based model for research.

The Exclusive Economic Zone of Chile defined by Easter Island and Salas y Gómez Island is in the South Pacific Sub-tropical Gyre (SPSG), putting it at the center of the most oligotrophic and biomass poor waters in the world. Only 10 biological oceanographic expeditions have entered this zone in 105 years (19052010). We review key aspects of the plankton ecosystem and biogeochemical function relevant for the understanding of and conservation planning for marine environments. Plankton production is limited by lack of dissolved inorganic fixed nitrogen, not phosphorous. Higher organic nitrogen levels might be biologically unavailable. Short-term experiments suggested iron is not limiting, yet iron still likely limits nitrogen fixation, and thus production, at longer time scales, as the presence of nitrogen-fixers is exceptionally low compared to other ocean gyres. Plankton function is dominated by the smallest unicellular organisms, picoplankton (

Following the publication of this article [1], the authors reported that the link to Additional file 11 linked to the wrong set of data. The correct supplementary data is provided in this Correction article (Additional file 11).

Adaptation to marginal habitats at species range-limits has often been associated with parthenogenetic reproduction in terrestrial animals and plants. Laboratory observations have shown that brown algae exhibit a high propensity for parthenogenesis by various mechanisms. The kelp Laminaria digitata is an important component of the ecosystem in Northern European rocky intertidal habitats. We studied four L. digitata populations for the effects of marginality on genetic diversity and sexual reproduction. Two populations were marginal: One (Locquirec, in Northern Brittany) was well within the geographic range, but was genetically isolated from other populations by large stretches of sandy beaches. Another population was at the range limits of the species (Quiberon, in Southern Brittany) and was exposed to much higher seasonal temperature changes. Microsatellite analyses confirmed that these populations showed decreased genetic and allelic diversity, consistent with marginality and genetic isolation. Sporophytes from both marginal populations showed greatly diminished spore-production compared to central populations, but only the southern-limit population (Quiberon) showed a high propensity for producing unreduced (2N) spores. Unreduced 2N spores formed phenotypically normal gametophytes with nuclear area consistent with ≥2N DNA contents, and microsatellite studies suggested these were produced at least in part by automixis. However, despite this being the dominant path of spore production in Quiberon sporophyte individuals, the genetic evidence indicated the population was maintained mostly by sexual reproduction. Thus, although spore production and development showed the expected tendency of geographical parthenogenesis in marginal populations, this appeared to be a consequence of maladaptation, rather than an adaptation to, life in a marginal habitat.

Successful completion of development requires coordination of patterning events with morphogenetic movements. Environmental variability challenges this coordination. For example, developing organisms encounter varying environmental temperatures that can strongly influence developmental rates. We hypothesized that the mechanics of morphogenesis would have to be finely adjusted to allow for normal morphogenesis across a wide range of developmental rates. We formulated our hypothesis as a simple model incorporating time-dependent application of force to a viscoelastic tissue. This model suggested that the capacity to maintain normal morphogenesis across a range of temperatures would depend on how both tissue viscoelasticity and the forces that drive deformation vary with temperature. To test this model we investigated how the mechanical behavior of embryonic tissue (Xenopus laevis) changed with temperature; we used a combination of micropipette aspiration to measure viscoelasticity, electrically induced contractions to measure cellular force generation, and confocal microscopy to measure endogenous contractility. Contrary to expectations, the viscoelasticity of the tissues and peak contractile tension proved invariant with temperature even as rates of force generation and gastrulation movements varied three-fold. Furthermore, the relative rates of different gastrulation movements varied with temperature: the speed of blastopore closure increased more slowly with temperature than the speed of the dorsal-to-ventral progression of involution. The changes in the relative rates of different tissue movements can be explained by the viscoelastic deformation model given observed viscoelastic properties, but only if morphogenetic forces increase slowly rather than all at once.

Previous studies suggest that mechanical feedback could coordinate morphogenetic events in embryos. Furthermore, embryonic tissues have complex structure and composition and undergo large deformations during morphogenesis. Hence we expect highly non-linear and loading-rate dependent tissue mechanical properties in embryos.We used micro-aspiration to test whether a simple linear viscoelastic model was sufficient to describe the mechanical behavior of gastrula stage Xenopus laevis embryonic tissue in vivo. We tested whether these embryonic tissues change their mechanical properties in response to mechanical stimuli but found no evidence of changes in the viscoelastic properties of the tissue in response to stress or stress application rate. We used this model to test hypotheses about the pattern of force generation during electrically induced tissue contractions. The dependence of contractions on suction pressure was most consistent with apical tension, and was inconsistent with isotropic contraction. Finally, stiffer clutches generated stronger contractions, suggesting that force generation and stiffness may be coupled in the embryo.The mechanical behavior of a complex, active embryonic tissue can be surprisingly well described by a simple linear viscoelastic model with power law creep compliance, even at high deformations. We found no evidence of mechanical feedback in this system. Together these results show that very simple mechanical models can be useful in describing embryo mechanics.

Abstract Background Diatoms are one of the most species-rich groups of eukaryotic microbes known. Diatoms are also the only group of eukaryotic micro-algae with a diplontic life history, suggesting that the ancestral diatom switched to a life history dominated by a duplicated genome. A key mechanism of speciation among diatoms could be a propensity for additional stable genome duplications. Across eukaryotic taxa, genome size is directly correlated to cell size and inversely correlated to physiological rates. Differences in relative genome size, cell size, and acclimated growth rates were analyzed in isolates of the diatom Ditylum brightwellii. Ditylum brightwellii consists of two main populations with identical 18s rDNA sequences; one population is distributed globally at temperate latitudes and the second appears to be localized to the Pacific Northwest coast of the USA. These two populations co-occur within the Puget Sound estuary of WA, USA, although their peak abundances differ depending on local conditions. Results All isolates from the more regionally-localized population (population 2) possessed 1.94 ± 0.74 times the amount of DNA, grew more slowly, and were generally larger than isolates from the more globally distributed population (population 1). The ITS1 sequences, cell sizes, and genome sizes of isolates from New Zealand were the same as population 1 isolates from Puget Sound, but their growth rates were within the range of the slower-growing population 2 isolates. Importantly, the observed genome size difference between isolates from the two populations was stable regardless of time in culture or the changes in cell size that accompany the diatom life history. Conclusions The observed two-fold difference in genome size between the D. brightwellii populations suggests that whole genome duplication occurred within cells of population 1 ultimately giving rise to population 2 cells. The apparent regional localization of population 2 is consistent with a recent divergence between the populations, which are likely cryptic species. Genome size variation is known to occur in other diatom genera; we hypothesize that genome duplication may be an active and important mechanism of genetic and physiological diversification and speciation in diatoms.