Mesocosm study of PAC-modified clay effects on Karenia brevis cells and toxins, chemical dynamics, and benthic invertebrate physiology.

Modified clay compounds are used globally as a method of controlling harmful algal blooms, and their use is currently under consideration to control Karenia brevis blooms in Florida, USA. In 1400 L mesocosm tanks, chemical dynamics and lethal and sublethal impacts of MC II, a polyaluminum chloride (PAC)-modified kaolinite clay, were evaluated over 72 h on a benthic community representative of Sarasota Bay, which included blue crab (Callinectes sapidus), sea urchin (Lytechinus variegatus), and hard clam (Mercenaria campechiensis). In this experiment, MC II was dosed at 0.2 g L-1 to treat bloom-level densities of K. brevis at 1 × 106 cells L-1. Cell removal in MC II-treated tanks was 57% after 8 h and 95% after 48 h. In the water column, brevetoxin analogs BTx-1 and BTx-2 were found to be significantly higher in untreated tanks at 24 and 48 h, while in MC II-treated tanks, BTx-3 was found to be higher at 48 h and BTx-B5 was found to be higher at 24 and 48 h. In MC II floc, we found no significant differences in BTx-1 or BTx-2 between treatments for any time point, while BTx-3 was found to be significantly higher in the MC II-treated tanks at 48 and 72 h, and BTx-B5 was higher in MC II-treated tanks at 24 and 72 h. Among various chemical dynamics observed, it was notable that dissolved phosphorus was consistently significantly lower in MC II tanks after 2 h, and that turbidity in MC II tanks returned to control levels 48 h after treatment. Dissolved inorganic carbon and total seawater alkalinity were significantly reduced in MC II tanks, and partial pressure of CO2 (pCO2) was significantly higher in the MC II-only treatment after 2 h. In MC II floc, particulate phosphorus was found to be significantly higher in MC II tanks after 24 h. In animals, lethal and sublethal responses to MC II-treated K. brevis did not differ from untreated K. brevis for either of our three species at any time point, suggesting MC II treatment at this dosage has negligible impacts to these species within 72 h of exposure. These results appear promising in terms of the environmental safety of MC II as a potential bloom control option, and we recommend scaling up MC II experiments to field trials in order to gain deeper understanding of MC II performance and dynamics in natural waters.

Exploring indirect effects of a classic trophic cascade between urchins and kelp on zooplankton and whales.

Kelp forest trophic cascades have been extensively researched, yet indirect effects to the zooplankton prey base and gray whales have not been explored. We investigate the correlative patterns of a trophic cascade between bull kelp and purple sea urchins on gray whales and zooplankton in Oregon, USA. Using generalized additive models (GAMs), we assess (1) temporal dynamics of the four species across 8 years, and (2) possible trophic paths from urchins to kelp, kelp as habitat to zooplankton, and kelp and zooplankton to gray whales. Temporal GAMs revealed an increase in urchin coverage, with simultaneous decline in kelp condition, zooplankton abundance and gray whale foraging time. Trophic path GAMs, which tested for correlations between species, demonstrated that urchins and kelp were negatively correlated, while kelp and zooplankton were positively correlated. Gray whales showed nuanced and site-specific correlations with zooplankton in one site, and positive correlations with kelp condition in both sites. The negative correlation between the kelp-urchin trophic cascade and zooplankton resulted in a reduced prey base for gray whales. This research provides a new perspective on the vital role kelp forests may play across multiple trophic levels and interspecies linkages.

Interactive effects of chronic ocean acidification and warming on the growth, survival, and physiological responses of adults of the temperate sea urchin Strongylocentrotusintermedius.

To investigate the interactive effects of chronic ocean acidification and warming (OAW) on the growth, survival, and physiological responses of sea urchins, adults of the temperate sea urchin Strongylocentrotus intermedius were incubated separately/jointly in acidic (ΔpHNBS = -0.5 units) and thermal (ΔT = +3.0 °C) seawater for 120 days under lab-controlled conditions based on the projected ocean pH and temperature for 2100 put forward by the Intergovernmental Panel on Climate Change (IPCC). Survival rate (SR), average food consumption rate (FCR), gut index (GuI), specific growth rate (SGR), digestive capability, energy production, and antioxidant capability were subsequently determined. The results showed that 1) the SR, FCR, GuI and SGR decreased sharply under OAW conditions. Significant interactive effects of OAW on SR and SGR were observed at 120 days post-incubation (dpi), and on FCR this occurred at 90 dpi. 2) OAW altered the activities of both digestive and antioxidant enzymes. There were significant interaction effects of OAW on the activities of amylase, trehalase, and superoxide dismutase. 3) The relative gene expression levels and activities of key enzymes involved in glycometabolism pathways (i.e., glycolysis and the tricarboxylic acid cycle) were significantly affected by OAW, resulting in an alteration of the total ATP content in the sea urchins. Interaction effects of OAW were observed in both relative gene expression and the activity of enzymes involved in glycolysis (hexokinase), the transformation of glycolysis end-products (lactate dehydrogenase), the tricarboxylic acid cycle (citrate synthetase), and ATP production (Na+/K+-ATPase). The data from this study will enrich our knowledge concerning the combined effects of global climate change on the survival, growth, and physiological responses of echinoderms.

Transgenerational acclimation to acidified seawater and gene expression patterns in a sea urchin.

Transgenerational responses of susceptible calcifying organisms to progressive ocean acidification are an important issue in reducing uncertainty of future predictions. In this study, a two-generation rearing experiment was conducted using mature Mesocentrotus nudus, a major edible sea urchin that occurs along the coasts of northern Japan. Morphological observations and comprehensive gene expression analysis (RNA-seq) of resulting larvae were performed to examine transgenerational acclimation to acidified seawater. Two generations of rearing experiments showed that larvae derived from parents acclimated to acidified seawater tended to have higher survival and show less reduction in body size when exposed to acidified seawater of the same pH, suggesting that a positive carry-over effect occurred. RNA-seq analysis showed that gene expression patterns of larvae originated from both acclimated and non-acclimated parents to acidified seawater tended to be different than control condition, and the gene expression pattern of larvae originated from acclimated parents was substantially different than that of larvae of non-acclimated and control parents.

Has the concept of polyspermy prevention been invented in the laboratory?

There is no evidence, nor need, for a fast block to polyspermy in animal oocytes. The idea that oocytes have evolved a mechanism to allow the entry of one spermatozoon and repel all others has, however, gained consensus over the last century. The main culprit is the sea urchin, which has been used for over a century in in vitro studies of the fertilization process. Images of sea urchin oocytes with thousands of sperm attached to the surface are commonplace in textbooks and appeal to the nature of the reader implying an intriguing surface mechanism of sperm selection despite these oocytes being fixed for photography (Figure ). The abundance of gametes in this marine invertebrate and the ease of experimentation have given us the possibility to elucidate many aspects of the mechanism of fertilization, but has also led to ongoing controversies in reproductive biology, one being polyspermy prevention. Kinetic experiments by Rothschild and colleagues in the 1950s led to the hypothesis of a fast partial block to polyspermy in sea urchin oocytes that reduced the probability of a second spermatozoon from entering the oocyte by 1/20th. In the 1970s, Jaffe and colleagues suggested, with circumstantial evidence, that this partial block was due to the sperm-induced depolarization of the oocyte plasma membrane. However, the fate of supernumerary spermatozoa is determined well before the plasma membrane of the oocyte depolarizes. Transmembrane voltage does not serve to regulate sperm entry. Scholastic texts have inadvertently promulgated this concept across the animal kingdom with no logical correlation or experimentation and, as of today, a molecular mechanism to regulate sperm entry in oocytes has not been identified.

Surviving in an Acidifying Ocean: Acid-Base Physiology and Energetics of the Sea Urchin Larva.

The sea urchin larva has been used by biologists for more than a century to study the development and evolution of animals. Surprisingly, very little information has been generated regarding the physiology of this small planktonic organism. However, in the context of anthropogenic CO2-driven ocean acidification (OA), the membrane transport physiology and energetics of this marine model organism have received considerable attention in the past decade. This has led to the discovery of new, exciting physiological systems, including a highly alkaline digestive tract and the calcifying primary mesenchyme cells that generate the larval skeleton. These physiological systems directly relate to the energetics of the organisms when challenged by OA. Here we review the latest membrane transport physiology and energetics in the sea urchin larva, we identify emerging questions, and we point to important future directions in the field of marine physiology in times of rapid climate change.

The behavior of sympatric sea urchin species across an ecosystem state gradient.

Background: In temperate macroalgal forests, sea urchins are considered as a keystone species due to their grazing ability. Given their potential to shape benthic communities, we monitored the habitat use by three sympatric sea urchin species and compared their behaviors in a vegetated habitat (VH) and an adjacent isoyake habitat (IH). Methods: We monitored the environmental conditions and sea urchin density along deep and shallow transects of the VH and IH for over a year. The benthic rugosity at both sites were also surveyed. A mark-recapture experiment was conducted on the two most abundant sea urchins, Diadema setosum and Heliocidaris crassispina, to elucidate sea urchin movement patterns and group dynamics. Results: We found that exposure to waves was highest at the VH while the IH was sheltered. The deep IH experienced the least amount of light due to high turbidity. Water temperature patterns were similar across sites. The VH benthic topography was more rugose compared to the smoother and silt-covered IH substate. Peak macroalgal bloom occurred three months earlier in IH, but macroalgae persisted longer at the shallow VH. Among the sympatric sea urchins, H. crassispina was most abundant at the shallow VH and was observed in pits and crevices. The most abundant across IH and in the deep VH was D. setosum, preferring either crevices or free-living, depending on hydrodynamic conditions. The least abundant species was D. savignyi, and most often observed in crevices. Small and medium sea urchins were most often observed at the IH site, whereas larger sea urchins were more likely observed at the VH. The mark-recapture study showed that D. setosum was found to displace further at the IH, and H. crassispina was more sedentary. Additionally, D. setosum was always observed in groups, whereas H. crassispina was always solitary. Discussion: The behaviors of sympatric urchins, Diadema savignyi, D. setosum and H. crassispina, differed in response to changes in the benthic environment and physical conditions. Sea urchin displacement increased when rugosity and wave action were low. Habitat preference shifted to crevices in seasons with high wave action. In general, the mark-recapture experiment showed that sea urchins displaced further at night.

Acclimation of the Antarctic sea urchin Sterechinus neumayeri to warmer temperatures involves a modulation of cellular machinery.

Global warming is threatening marine Antarctic fauna, which has evolved in isolation in a cold environment for millions of years. Facing increasing temperatures, marine Antarctic invertebrates can either tolerate or develop adaptations to these changes. On a short timescale, their survival and resistance to warming will be driven by the efficiency of their phenotypic plasticity through their capacity for acclimation. The current study aims at evaluating the capacity for acclimation of the Antarctic sea urchin Sterechinus neumayeri to predicted ocean warming scenarios (+2, RCP 2.6 and + 4 °C, RCP 8.5, IPCC et al., 2019) and deciphering the subcellular mechanisms underlying their acclimation. A combination of transcriptomics, physiological (e.g. growth rate, gonad growth, ingestion rate and oxygen consumption), and behavioral-based approaches were used on individuals incubated at 1, 3 and, 5 °C for 22 weeks. Mortality was low at warmer temperatures (20%) and oxygen consumption and ingestion rate seemed to reach a stable state around 16 weeks suggesting that S. neumayeri might be able to acclimate to warmer temperatures (until 5 °C). Transcriptomic analyses highlighted adjustments of the cellular machinery with the activation of replication, recombination, and repair processes as well as cell cycle and division and repression of transcriptional and signal transduction mechanisms and defense processes. These results suggest that acclimation to warmer scenarios might require more than 22 weeks for the Antarctic Sea urchins S. neumayeri but that projections of climate change for the end of the century may not strongly affect the population of S. neumayeri of this part of the Antarctic.

Sunflower sea star predation on urchins can facilitate kelp forest recovery.

The recent collapse of predatory sunflower sea stars (Pycnopodia helianthoides) owing to sea star wasting disease (SSWD) is hypothesized to have contributed to proliferation of sea urchin barrens and losses of kelp forests on the North American west coast. We used experiments and a model to test whether restored Pycnopodia populations may help recover kelp forests through their consumption of nutritionally poor purple sea urchins (Strongylocentrotus purpuratus) typical of barrens. Pycnopodia consumed 0.68 S. purpuratus d-1, and our model and sensitivity analysis shows that the magnitude of recent Pycnopodia declines is consistent with urchin proliferation after modest sea urchin recruitment, and even small Pycnopodia recoveries could generally lead to lower densities of sea urchins that are consistent with kelp-urchin coexistence. Pycnopodia seem unable to chemically distinguish starved from fed urchins and indeed have higher predation rates on starved urchins owing to shorter handling times. These results highlight the importance of Pycnopodia in regulating purple sea urchin populations and maintaining healthy kelp forests through top-down control. The recovery of this important predator to densities commonly found prior to SSWD, whether through natural means or human-assisted reintroductions, may therefore be a key step in kelp forest restoration at ecologically significant scales.

Conspecific cues, not starvation, mediate barren urchin response to predation risk.

Prey state and prey density mediate antipredator responses that can shift community structure and alter ecosystem processes. For example, well-nourished prey at low densities (i.e., prey with higher per capita predation risk) should respond strongly to predators. Although prey state and density often co-vary across habitats, it is unclear if prey responses to predator cues are habitat-specific. We used mesocosms to compare the habitat-specific responses of purple sea urchins (Strongylocentrotus purpuratus) to waterborne cues from predatory lobsters (Panulirus interruptus). We predicted that urchins from kelp forests (i.e., in well-nourished condition) tested at low densities typically observed in this habitat would respond more strongly to predation risk than barren urchins (i.e., in less nourished condition) tested at high densities typically observed in this habitat. Indeed, when tested at densities associated with respective habitats, urchins from forests, but not barrens, reduced kelp grazing by 69% when exposed to lobster risk cues. Barren urchins that were unresponsive to predator cues at natural, high densities suddenly responded strongly to lobster cues when conspecific densities were reduced. Strong responses of low densities of barren urchins persisted across feeding history (i.e. 0-64 days of starvation). This suggests that barren urchins can respond to predators but typically do not because of high conspecific densities. Because high densities of urchins in barrens should weaken the non-consumptive effects of lobsters, urchins in these habitats may continue to graze in the presence of predators thereby providing a feedback that maintains urchin barrens.

Reference Genome of the California Sheephead, Semicossyphus pulcher (Labridae, Perciformes), A Keystone Fish Predator in Kelp Forest Ecosystems.

Keystone species are known to play a critical role in kelp forest health, including the well-known killer whales, sea otter, sea urchin, kelp trophic cascade in the Aleutian Islands, Alaska, USA. In California, a major player in the regulation of sea urchin abundance, and in turn, the health of kelp forests ecosystems, is a large wrasse, the California Sheephead, Semicossyphus pulcher. We present a reference genome for this ecologically important species that will serve as a key resource for future conservation research of California's inshore marine environment utilizing genomic tools to address changes in life-history traits, dispersal, range shifts, and ecological interactions among members of the kelp forest ecological assemblages. Our genome assembly of S. pulcher has a total length of 0.794 Gb, which is similar to many other marine fishes. The assembly is largely contiguous (N50 = 31.9 Mb) and nearly complete (BUSCO single-copy core gene content = 98.1%). Within the context of the California Conservation Genomics Project (CCGP), the genome of S. pulcher will be used as an important reference resource for ongoing whole genome resequencing efforts of the species.

Ocean acidification impacts sperm swimming performance and pHi in the New Zealand sea urchin Evechinus chloroticus.

In sea urchins, spermatozoa are stored in the gonads in hypercapnic conditions (pH<7.0). During spawning, sperm are diluted in seawater of pH>8.0, and there is an alkalinization of the sperm's internal pH (pHi) through the release of CO2 and H+. Previous research has shown that when pHi is above 7.2-7.3, the dynein ATPase flagellar motors are activated, and the sperm become motile. It has been hypothesized that ocean acidification (OA), which decreases the pH of seawater, may have a narcotic effect on sea urchin sperm by impairing the ability to regulate pHi, resulting in decreased motility and swimming speed. Here, we used data collected from the same individuals to test the relationship between pHi and sperm motility/performance in the New Zealand sea urchin Evechinus chloroticus under near-future (2100) and far-future (2150) atmospheric PCO2 conditions (RCP 8.5: pH 7.77, 7.51). Decreasing seawater pH significantly negatively impacted the proportion of motile sperm, and four of the six computer-assisted sperm analysis (CASA) sperm performance measures. In control conditions, sperm had an activated pHi of 7.52. Evechinus chloroticus sperm could not defend pHi in future OA conditions; there was a stepped decrease in the pHi at pH 7.77, with no significant difference in mean pHi between pH 7.77 and 7.51. Paired measurements in the same males showed a positive relationship between pHi and sperm motility, but with a significant difference in the response between males. Differences in motility and sperm performance in OA conditions may impact fertilization success in a future ocean.

Alternations in the foraging behaviour of a primary consumer drive patch transition dynamics in a temperate rocky reef ecosystem.

Understanding the role of animal behaviour in linking individuals to ecosystems is central to advancing knowledge surrounding community structure, stability and transition dynamics. Using 22 years of long-term subtidal monitoring, we show that an abrupt outbreak of purple sea urchins (Strongylocentrotus purpuratus), which occurred in 2014 in southern Monterey Bay, California, USA, was primarily driven by a behavioural shift, not by a demographic response (i.e. survival or recruitment). We then tracked the foraging behaviour of sea urchins for 3 years following the 2014 outbreak and found that behaviour is strongly associated with patch state (forest or barren) transition dynamics. Finally, in 2019, we observed a remarkable recovery of kelp forests at a deep rocky reef. We show that this recovery was associated with sea urchin movement from the deep reef to shallow water. These results demonstrate how changes in grazer behaviour can facilitate patch dynamics and dramatically restructure communities and ecosystems.

Macroalgae and interspecific alarm cues regulate behavioral interactions between sea urchins and sea cucumbers.

Sea urchins and sea cucumbers are mutually beneficial organisms in kelp ecosystem. As herbivores, sea urchins process kelp through feeding and egestion, providing inaccessible food for benthic consumers such as sea cucumbers. Sea urchins in turn profit from the sediment cleaned by sea cucumbers. However, behavioral interactions between them remain poorly understood, which greatly hampers our understanding on the relationship between ecologically important benthic species in marine ecosystems and the regulating mechanism. The present study investigated behavioral interactions between sea urchins Strongylocentrotus intermedius and sea cucumbers Apostichopus japonicus in laboratory conditions. We revealed that the presence of sea urchins caused significant higher speed movement of A. japonicus. Interestingly, the negative effects of S. intermedius on A. japonicus were significantly reduced in the shared macroalgal area. For the first time, we found the interspecific responses to alarm cues between sea cucumbers and sea urchins. Conspecific responses were significantly larger than the interspecific responses in both sea urchins and sea cucumbers. This indicates that interspecific response to alarm cues is an efficient approach to anti-predation and coexistence in mutually beneficial organisms. The present study shed light on the interspecific relationships and coexistence between sea urchins and sea cucumbers in kelp ecosystem.

Global kelp forest restoration: past lessons, present status, and future directions.

Kelp forest ecosystems and their associated ecosystem services are declining around the world. In response, marine managers are working to restore and counteract these declines. Kelp restoration first started in the 1700s in Japan and since then has spread across the globe. Restoration efforts, however, have been largely disconnected, with varying methodologies trialled by different actors in different countries. Moreover, a small subset of these efforts are 'afforestation', which focuses on creating new kelp habitat, as opposed to restoring kelp where it previously existed. To distil lessons learned over the last 300 years of kelp restoration, we review the history of kelp restoration (including afforestation) around the world and synthesise the results of 259 documented restoration attempts spanning from 1957 to 2020, across 16 countries, five languages, and multiple user groups. Our results show that kelp restoration projects have increased in frequency, have employed 10 different methodologies and targeted 17 different kelp genera. Of these projects, the majority have been led by academics (62%), have been conducted at sizes of less than 1 ha (80%) and took place over time spans of less than 2 years. We show that projects are most successful when they are located near existing kelp forests. Further, disturbance events such as sea-urchin grazing are identified as regular causes of project failure. Costs for restoration are historically high, averaging hundreds of thousands of dollars per hectare, therefore we explore avenues to reduce these costs and suggest financial and legal pathways for scaling up future restoration efforts. One key suggestion is the creation of a living database which serves as a platform for recording restoration projects, showcasing and/or re-analysing existing data, and providing updated information. Our work establishes the groundwork to provide adaptive and relevant recommendations on best practices for kelp restoration projects today and into the future.

Staying in place and moving in space: Contrasting larval thermal sensitivity explains distributional changes of sympatric sea urchin species to habitat warming.

For marine ectotherms, larval success, planktonic larval duration and dispersal trajectories are strongly influenced by temperature, and therefore, ocean warming and heatwaves have profound impacts on these sensitive stages. Warming, through increased poleward flow in regions with western boundary currents, such as the East Australia Current (EAC), provides opportunities for range extension as propagules track preferred conditions. Two sea urchin species, Centrostephanus rodgersii and Heliocidaris tuberculata, sympatric in the EAC warming hotspot, exhibit contrasting responses to warming. Over half a century, C. rodgersii has undergone marked poleward range extension, but the range of H. tuberculata has not changed. We constructed thermal performance curves (TPC) to determine if contrasting developmental thermal tolerance can explain this difference. The temperatures tested encompassed present-day distribution and forecast ocean warming/heatwave conditions. The broad and narrow thermal optimum (Topt) ranges for C. rodgersii and H. tuberculata larvae (7.2 and 4.7°C range, respectively) matched their realized (adult distribution) thermal niches. The cool and warm temperatures for 50% development to the feeding larva approximated temperatures at adult poleward range limits. Larval cool tolerances with respect to mean local temperature differed, 6.0 and 3.8°C respectively. Larval warm tolerances were similar for both species as are the adult warm range edges. The larvae of both species would be sensitive to heatwaves. Centrostephanus rodgersii has stayed in place and shifted in space, likely due to its broad cold-warm larval thermal tolerance and large thermal safety margins. Phenotypic plasticity of the planktonic stage of C. rodgersii facilitated its range extension. In contrast, larval cold intolerance of H. tuberculata explains its restricted range and will delay poleward extension as the region warms. In a warming ocean, we show that intrinsic thermal biology traits of the pelagic stage provide an integrative tool to explain species-specific variation in range shift patterns.

Interactive effects of ocean acidification, ocean warming, and diurnal temperature cycling on antioxidant responses and energy budgets in two sea urchins Strongylocentrotus intermedius and Tripneustes gratilla from different latitudes.

To accurately predict the fitness of marine ectotherms under the climate change scenarios, interactive effects from multiple environmental stressors should be considered, such as ocean acidification (OA), ocean warming (OW) and diurnal temperature cycling (DTC). In this work, we evaluated and compared the antioxidant capacity and metabolism homeostasis of two sea urchins, viz. the temperate species Strongylocentrotus intermedius and the tropical species Tripneustes gratilla, in response to oceanic conditions under a climate change scenario. The two species were treated separately/jointly by acidic (pH 7.6), thermal (ambient temperature + 3 °C), and temperature fluctuating (5 °C fluctuations daily) seawater for 28 days. The activities of antioxidant enzymes (catalase and superoxide dismutase) and the cellular energy allocation in the urchins' gonads were assessed subsequently. Results showed that exposure to OA, OW, and DTC all induced antioxidant responses associated with metabolism imbalance in both S. intermedius and T. gratilla. The physiological adjustments and energy strategies towards exposure of OA, OW, and DTC are species specific, perhaps owing to the different thermal acclimation of species from two latitudes. Moreover, decrease of cellular energy allocation were detected in both species under combined OA, OW, and DTC conditions, indicating unsustainable bioenergetic states. The decrease of cellular energy allocation is weaker in T. gratilla than in S. intermedius, implying higher acclimation capacity to maintain the energy homeostasis in tropical urchins. These results suggest that climate change might affect the population replenishment of the two sea urchins species, especially for the temperate species.

High fitness areas drive the aggregation of the sea urchin Mesocentrotus nudus.

Sea urchin aggregation is a common phenomenon in coastlines. However, it remains controversial whether sea urchins form resource aggregations or behavioral aggregations in a non-spawning season. To clarify, we studied the aggregative responses to food and predators in the sea urchin Mesocentrotus nudus when high fitness areas (HFAs) were scarce versus sufficient. By taking the occupied area of each sea urchin (test diameter + spines =  4.5 cm) as a square (4.5 cm × 4.5 cm), we set scarce HFAs for the sea urchins in Experiment 1 (the squares of HFAs: the area occupied by experimental sea urchins = 1:1) and sufficient HFAs for the sea urchins in Experiment 2 (the squares of HFAs: the area occupied by experimental sea urchins = 2:1). If M. nudus form resource aggregations, they would aggregate passively under the scarce HFAs conditions, but not in the sufficient HFAs conditions. Conversely, if M. nudus form behavioral aggregation, aggregation would occur in both scarce and sufficient HFAs. The present results showed that in the scarce HFAs, M. nudus in the food and predator groups were significantly closer to the food and further from predators, and had significantly more aggregated numbers in HFAs than those in the control group. Sea urchins did not aggregate in response to food or predators under the sufficient HFAs, although significantly more sea urchins of the experimental group was found in HFAs than that of the control group. Sea urchins (at least M. nudus) form resource aggregations that are driven by the scarce HFAs. This provides valuable information into the mechanisms of the aggregation of sea urchins.

Plasticity in fluctuating hydrodynamic conditions: tube foot regeneration in sea urchins.

Regenerating structures critical for survival provide excellent model systems for the study of phenotypic plasticity. These body components must regenerate their morphology and functionality quickly while subjected to different environmental stressors. Sea urchins live in high-energy environments where hydrodynamic conditions pose significant challenges. Adhesive tube feet provide secure attachment to the substratum but can be amputated by predation and hydrodynamic forces. Tube feet display functional and morphological plasticity in response to environmental conditions, but regeneration to their pre-amputation status has not been achieved under quiescent laboratory settings. In this study, we assessed the effect of turbulent water movement, periodic emersion and quiescent conditions on the regeneration process of tube foot morphology (length, disc area) and functionality (maximum disc tenacity, stem breaking force). Disc area showed significant plasticity in response to the treatments; when exposed to emersion and turbulent water movement, disc area was larger than that of tube feet regenerated in quiescent conditions. However, no treatment stimulated regeneration to pre-amputation sizes. Tube foot length was unaffected by treatments and remained shorter than non-amputated tube feet. Stem breaking force for amputated and non-amputated treatments increased in all cases when compared with pre-amputation values. Maximum tenacity (force per unit area) was similar among tube feet subjected to simulated field conditions and amputation treatments. Our results suggest a role of active plasticity of tube foot functional morphology in response to field-like conditions and demonstrate the plastic response of invertebrates to laboratory conditions.