Global warming impairs stock-recruitment dynamics of corals.

Changes in disturbance regimes due to climate change are increasingly challenging the capacity of ecosystems to absorb recurrent shocks and reassemble afterwards, escalating the risk of widespread ecological collapse of current ecosystems and the emergence of novel assemblages1-3. In marine systems, the production of larvae and recruitment of functionally important species are fundamental processes for rebuilding depleted adult populations, maintaining resilience and avoiding regime shifts in the face of rising environmental pressures4,5. Here we document a regional-scale shift in stock-recruitment relationships of corals along the Great Barrier Reef-the world's largest coral reef system-following unprecedented back-to-back mass bleaching events caused by global warming. As a consequence of mass mortality of adult brood stock in 2016 and 2017 owing to heat stress6, the amount of larval recruitment declined in 2018 by 89% compared to historical levels. For the first time, brooding pocilloporids replaced spawning acroporids as the dominant taxon in the depleted recruitment pool. The collapse in stock-recruitment relationships indicates that the low resistance of adult brood stocks to repeated episodes of coral bleaching is inexorably tied to an impaired capacity for recovery, which highlights the multifaceted processes that underlie the global decline of coral reefs. The extent to which the Great Barrier Reef will be able to recover from the collapse in stock-recruitment relationships remains uncertain, given the projected increased frequency of extreme climate events over the next two decades7.

Promoting inclusive metrics of success and impact to dismantle a discriminatory reward system in science.

Success and impact metrics in science are based on a system that perpetuates sexist and racist "rewards" by prioritizing citations and impact factors. These metrics are flawed and biased against already marginalized groups and fail to accurately capture the breadth of individuals' meaningful scientific impacts. We advocate shifting this outdated value system to advance science through principles of justice, equity, diversity, and inclusion. We outline pathways for a paradigm shift in scientific values based on multidimensional mentorship and promoting mentee well-being. These actions will require collective efforts supported by academic leaders and administrators to drive essential systemic change.

Evolved differences in energy metabolism and growth dictate the impacts of ocean acidification on abalone aquaculture.

Ocean acidification (OA) poses a major threat to marine ecosystems and shellfish aquaculture. A promising mitigation strategy is the identification and breeding of shellfish varieties exhibiting resilience to acidification stress. We experimentally compared the effects of OA on two populations of red abalone (Haliotis rufescens), a marine mollusc important to fisheries and global aquaculture. Results from our experiments simulating captive aquaculture conditions demonstrated that abalone sourced from a strong upwelling region were tolerant of ongoing OA, whereas a captive-raised population sourced from a region of weaker upwelling exhibited significant mortality and vulnerability to OA. This difference was linked to population-specific variation in the maternal provisioning of lipids to offspring, with a positive correlation between lipid concentrations and survival under OA. This relationship also persisted in experiments on second-generation animals, and larval lipid consumption rates varied among paternal crosses, which is consistent with the presence of genetic variation for physiological traits relevant for OA survival. Across experimental trials, growth rates differed among family lineages, and the highest mortality under OA occurred in the fastest growing crosses. Identifying traits that convey resilience to OA is critical to the continued success of abalone and other shellfish production, and these mitigation efforts should be incorporated into breeding programs for commercial and restoration aquaculture.

Characterization of Upper Extremity Kinematics Using Virtual Reality Movement Tasks and Wearable IMU Technology.

Task-specific training has been shown to be an effective neuromotor rehabilitation intervention, however, this repetitive approach is not always very engaging. Virtual reality (VR) systems are becoming increasingly popular in therapy due to their ability to encourage movement through customizable and immersive environments. Additionally, VR can allow for a standardization of tasks that is often lacking in upper extremity research. Here, 16 healthy participants performed upper extremity movement tasks synced to music, using a commercially available VR game known as Beat Saber. VR tasks were customized to characterize participants' joint angles with respect to each task's specified cardinal direction (inward, outward, upward, or downward) and relative task location (medial, lateral, high, and/or low). Movement levels were designed using three common therapeutic approaches: (1) one arm moving only (unilateral), (2) two arms moving in mirrored directions about the participant's midline (mirrored), or (3) two arms moving in opposing directions about the participant's midline (opposing). Movement was quantified using an XSens System, a wearable inertial measurement unit (IMU) technology. Results reveal a highly engaging and effective approach to quantifying movement strategies. Inward and outward (horizontal) tasks resulted in decreased wrist extension. Upward and downward (vertical) tasks resulted in increased shoulder flexion, wrist radial deviation, wrist ulnar deviation, and elbow flexion. Lastly, compared to opposing, mirrored, and unilateral movement levels often exaggerated joint angles. Virtual reality games, like Beat Saber, offer a repeatable and customizable upper extremity intervention that has the potential to increase motivation in therapeutic applications.

Coast-wide evidence of low pH amelioration by seagrass ecosystems.

Global-scale ocean acidification has spurred interest in the capacity of seagrass ecosystems to increase seawater pH within crucial shoreline habitats through photosynthetic activity. However, the dynamic variability of the coastal carbonate system has impeded generalization into whether seagrass aerobic metabolism ameliorates low pH on physiologically and ecologically relevant timescales. Here we present results of the most extensive study to date of pH modulation by seagrasses, spanning seven meadows (Zostera marina) and 1000 km of U.S. west coast over 6 years. Amelioration by seagrass ecosystems compared to non-vegetated areas occurred 65% of the time (mean increase 0.07 ± 0.008 SE). Events of continuous elevation in pH within seagrass ecosystems, indicating amelioration of low pH, were longer and of greater magnitude than opposing cases of reduced pH or exacerbation. Sustained elevations in pH of >0.1, comparable to a 30% decrease in [H+ ], were not restricted only to daylight hours but instead persisted for up to 21 days. Maximal pH elevations occurred in spring and summer during the seagrass growth season, with a tendency for stronger effects in higher latitude meadows. These results indicate that seagrass meadows can locally alleviate low pH conditions for extended periods of time with important implications for the conservation and management of coastal ecosystems.

Seagrass-driven changes in carbonate chemistry enhance oyster shell growth.

Quantifying the strength of non-trophic interactions exerted by foundation species is critical to understanding how natural communities respond to environmental stress. In the case of ocean acidification (OA), submerged marine macrophytes, such as seagrasses, may create local areas of elevated pH due to their capacity to sequester dissolved inorganic carbon through photosynthesis. However, although seagrasses may increase seawater pH during the day, they can also decrease pH at night due to respiration. Therefore, it remains unclear how consequences of such diel fluctuations may unfold for organisms vulnerable to OA. We established mesocosms containing different levels of seagrass biomass (Zostera marina) to create a gradient of carbonate chemistry conditions and explored consequences for growth of juvenile and adult oysters (Crassostrea gigas), a non-native species widely used in aquaculture that can co-occur, and is often grown, in proximity to seagrass beds. In particular, we investigated whether increased diel fluctuations in pH due to seagrass metabolism affected oyster growth. Seagrasses increased daytime pH up to 0.4 units but had little effect on nighttime pH (reductions less than 0.02 units). Thus, both the average pH and the amplitude of diel pH fluctuations increased with greater seagrass biomass. The highest seagrass biomass increased oyster shell growth rate (mm day-1) up to 40%. Oyster somatic tissue weight and oyster condition index exhibited a different pattern, peaking at intermediate levels of seagrass biomass. This work demonstrates the ability of seagrasses to facilitate oyster calcification and illustrates how non-trophic metabolic interactions can modulate effects of environmental change.

Response of seafloor ecosystems to abrupt global climate change.

Anthropogenic climate change is predicted to decrease oceanic oxygen (O2) concentrations, with potentially significant effects on marine ecosystems. Geologically recent episodes of abrupt climatic warming provide opportunities to assess the effects of changing oxygenation on marine communities. Thus far, this knowledge has been largely restricted to investigations using Foraminifera, with little being known about ecosystem-scale responses to abrupt, climate-forced deoxygenation. We here present high-resolution records based on the first comprehensive quantitative analysis, to our knowledge, of changes in marine metazoans (Mollusca, Echinodermata, Arthropoda, and Annelida; >5,400 fossils and trace fossils) in response to the global warming associated with the last glacial to interglacial episode. The molluscan archive is dominated by extremophile taxa, including those containing endosymbiotic sulfur-oxidizing bacteria (Lucinoma aequizonatum) and those that graze on filamentous sulfur-oxidizing benthic bacterial mats (Alia permodesta). This record, from 16,100 to 3,400 y ago, demonstrates that seafloor invertebrate communities are subject to major turnover in response to relatively minor inferred changes in oxygenation (>1.5 to <0.5 mL⋅L(-1) [O2]) associated with abrupt (<100 y) warming of the eastern Pacific. The biotic turnover and recovery events within the record expand known rates of marine biological recovery by an order of magnitude, from <100 to >1,000 y, and illustrate the crucial role of climate and oceanographic change in driving long-term successional changes in ocean ecosystems.

Interactive effects of temperature, food and skeletal mineralogy mediate biological responses to ocean acidification in a widely distributed bryozoan.

Marine invertebrates with skeletons made of high-magnesium calcite may be especially susceptible to ocean acidification (OA) due to the elevated solubility of this form of calcium carbonate. However, skeletal composition can vary plastically within some species, and it is largely unknown how concurrent changes in multiple oceanographic parameters will interact to affect skeletal mineralogy, growth and vulnerability to future OA. We explored these interactive effects by culturing genetic clones of the bryozoan Jellyella tuberculata (formerly Membranipora tuberculata) under factorial combinations of dissolved carbon dioxide (CO2), temperature and food concentrations. High CO2 and cold temperature induced degeneration of zooids in colonies. However, colonies still maintained high growth efficiencies under these adverse conditions, indicating a compensatory trade-off whereby colonies degenerate more zooids under stress, redirecting energy to the growth and maintenance of new zooids. Low-food concentration and elevated temperatures also had interactive effects on skeletal mineralogy, resulting in skeletal calcite with higher concentrations of magnesium, which readily dissolved under high CO2 For taxa that weakly regulate skeletal magnesium concentration, skeletal dissolution may be a more widespread phenomenon than is currently documented and is a growing concern as oceans continue to warm and acidify.

Plastic responses of bryozoans to ocean acidification.

Phenotypic plasticity has the potential to allow organisms to respond rapidly to global environmental change, but the range and effectiveness of these responses are poorly understood across taxa and growth strategies. Colonial organisms might be particularly resilient to environmental stressors, as organizational modularity and successive asexual generations can allow for distinctively flexible responses in the aggregate form. We performed laboratory experiments to examine the effects of increasing dissolved carbon dioxide (CO2) (i.e. ocean acidification) on the colonial bryozoan Celleporella cornuta sampled from two source populations within a coastal upwelling region of the northern California coast. Bryozoan colonies were remarkably plastic under these CO2 treatments. Colonies raised under high CO2 grew more quickly, investing less in reproduction and producing lighter skeletons when compared with genetically identical clones raised under current surface atmosphere CO2 values. Bryozoans held under high CO2 conditions also changed the Mg/Ca ratio of skeletal calcite and increased the expression of organic coverings in new growth, which may serve as protection against acidified water. We also observed strong differences between source populations in reproductive investment and organic covering reaction norms, consistent with adaptive responses to persistent spatial variation in local oceanographic conditions. Our results demonstrate that phenotypic plasticity and energetic trade-offs can mediate biological responses to global environmental change, and highlight the broad range of strategies available to colonial organisms.

Interacting environmental mosaics drive geographic variation in mussel performance and predation vulnerability.

Although theory suggests geographic variation in species' performance is determined by multiple niche parameters, little consideration has been given to the spatial structure of interacting stressors that may shape local and regional vulnerability to global change. Here, we use spatially explicit mosaics of carbonate chemistry, food availability and temperature spanning 1280 km of coastline to test whether persistent, overlapping environmental mosaics mediate the growth and predation vulnerability of a critical foundation species, the mussel Mytilus californianus. We find growth was highest and predation vulnerability was lowest in dynamic environments with frequent exposure to low pH seawater and consistent food. In contrast, growth was lowest and predation vulnerability highest when exposure to low pH seawater was decoupled from high food availability, or in exceptionally warm locations. These results illustrate how interactions among multiple drivers can cause unexpected, yet persistent geographic mosaics of species performance, interactions and vulnerability to environmental change.

Ocean acidification alters the response of intertidal snails to a key sea star predator.

Organism-level effects of ocean acidification (OA) are well recognized. Less understood are OA's consequences for ecological species interactions. Here, we examine a behaviourally mediated predator-prey interaction within the rocky intertidal zone of the temperate eastern Pacific Ocean, using it as a model system to explore OA's capacity to impair invertebrate anti-predator behaviours more broadly. Our system involves the iconic sea star predator, Pisaster ochraceus, that elicits flee responses in numerous gastropod prey. We examine, in particular, the capacity for OA-associated reductions in pH to alter flight behaviours of the black turban snail, Tegula funebralis, an often-abundant and well-studied grazer in the system. We assess interactions between these species at 16 discrete levels of pH, quantifying the full functional response of Tegula under present and near-future OA conditions. Results demonstrate the disruption of snail anti-predator behaviours at low pH, with decreases in the time individuals spend in refuge locations. We also show that fluctuations in pH, including those typical of rock pools inhabited by snails, do not materially change outcomes, implying little capacity for episodically benign pH conditions to aid behavioural recovery. Together, these findings suggest a strong potential for OA to induce cascading community-level shifts within this long-studied ecosystem.

Upper-ocean-to-atmosphere radiocarbon offsets imply fast deglacial carbon dioxide release.

Radiocarbon in the atmosphere is regulated largely by ocean circulation, which controls the sequestration of carbon dioxide (CO(2)) in the deep sea through atmosphere-ocean carbon exchange. During the last glaciation, lower atmospheric CO(2) levels were accompanied by increased atmospheric radiocarbon concentrations that have been attributed to greater storage of CO(2) in a poorly ventilated abyssal ocean. The end of the ice age was marked by a rapid increase in atmospheric CO(2) concentrations that coincided with reduced (14)C/(12)C ratios (Delta(14)C) in the atmosphere, suggesting the release of very 'old' ((14)C-depleted) CO(2) from the deep ocean to the atmosphere. Here we present radiocarbon records of surface and intermediate-depth waters from two sediment cores in the southwest Pacific and Southern oceans. We find a steady 170 per mil decrease in Delta(14)C that precedes and roughly equals in magnitude the decrease in the atmospheric radiocarbon signal during the early stages of the glacial-interglacial climatic transition. The atmospheric decrease in the radiocarbon signal coincides with regionally intensified upwelling and marine biological productivity, suggesting that CO(2) released by means of deep water upwelling in the Southern Ocean lost most of its original depleted-(14)C imprint as a result of exchange and isotopic equilibration with the atmosphere. Our data imply that the deglacial (14)C depletion previously identified in the eastern tropical North Pacific must have involved contributions from sources other than the previously suggested carbon release by way of a deep Southern Ocean pathway, and may reflect the expanded influence of the (14)C-depleted North Pacific carbon reservoir across this interval. Accordingly, shallow water masses advecting north across the South Pacific in the early deglaciation had little or no residual (14)C-depleted signals owing to degassing of CO(2) and biological uptake in the Southern Ocean.

Evolutionary change during experimental ocean acidification.

Rising atmospheric carbon dioxide (CO2) conditions are driving unprecedented changes in seawater chemistry, resulting in reduced pH and carbonate ion concentrations in the Earth's oceans. This ocean acidification has negative but variable impacts on individual performance in many marine species. However, little is known about the adaptive capacity of species to respond to an acidified ocean, and, as a result, predictions regarding future ecosystem responses remain incomplete. Here we demonstrate that ocean acidification generates striking patterns of genome-wide selection in purple sea urchins (Strongylocentrotus purpuratus) cultured under different CO2 levels. We examined genetic change at 19,493 loci in larvae from seven adult populations cultured under realistic future CO2 levels. Although larval development and morphology showed little response to elevated CO2, we found substantial allelic change in 40 functional classes of proteins involving hundreds of loci. Pronounced genetic changes, including excess amino acid replacements, were detected in all populations and occurred in genes for biomineralization, lipid metabolism, and ion homeostasis--gene classes that build skeletons and interact in pH regulation. Such genetic change represents a neglected and important impact of ocean acidification that may influence populations that show few outward signs of response to acidification. Our results demonstrate the capacity for rapid evolution in the face of ocean acidification and show that standing genetic variation could be a reservoir of resilience to climate change in this coastal upwelling ecosystem. However, effective response to strong natural selection demands large population sizes and may be limited in species impacted by other environmental stressors.

Ocean acidification research in the 'post-genomic' era: Roadmaps from the purple sea urchin Strongylocentrotus purpuratus.

Advances in nucleic acid sequencing technology are removing obstacles that historically prevented use of genomics within ocean change biology. As one of the first marine calcifiers to have its genome sequenced, purple sea urchins (Strongylocentrotus purpuratus) have been the subject of early research exploring genomic responses to ocean acidification, work that points to future experiments and illustrates the value of expanding genomic resources to other marine organisms in this new 'post-genomic' era. This review presents case studies of S. purpuratus demonstrating the ability of genomic experiments to address major knowledge gaps within ocean acidification. Ocean acidification research has focused largely on species vulnerability, and studies exploring mechanistic bases of tolerance toward low pH seawater are comparatively few. Transcriptomic responses to high pCO2 seawater in a population of urchins already encountering low pH conditions have cast light on traits required for success in future oceans. Secondly, there is relatively little information on whether marine organisms possess the capacity to adapt to oceans progressively decreasing in pH. Genomics offers powerful methods to investigate evolutionary responses to ocean acidification and recent work in S. purpuratus has identified genes under selection in acidified seawater. Finally, relatively few ocean acidification experiments investigate how shifts in seawater pH combine with other environmental factors to influence organism performance. In S. purpuratus, transcriptomics has provided insight into physiological responses of urchins exposed simultaneously to warmer and more acidic seawater. Collectively, these data support that similar breakthroughs will occur as genomic resources are developed for other marine species.

Ocean acidification increases the vulnerability of native oysters to predation by invasive snails.

There is growing concern that global environmental change might exacerbate the ecological impacts of invasive species by increasing their per capita effects on native species. However, the mechanisms underlying such shifts in interaction strength are poorly understood. Here, we test whether ocean acidification, driven by elevated seawater pCO2, increases the susceptibility of native Olympia oysters to predation by invasive snails. Oysters raised under elevated pCO2 experienced a 20% increase in drilling predation. When presented alongside control oysters in a choice experiment, 48% more high-CO2 oysters were consumed. The invasive snails were tolerant of elevated CO2 with no change in feeding behaviour. Oysters raised under acidified conditions did not have thinner shells, but were 29-40% smaller than control oysters, and these smaller individuals were consumed at disproportionately greater rates. Reduction in prey size is a common response to environmental stress that may drive increasing per capita effects of stress-tolerant invasive predators.

Larval carry-over effects from ocean acidification persist in the natural environment.

An extensive body of work suggests that altered marine carbonate chemistry can negatively influence marine invertebrates, but few studies have examined how effects are moderated and persist in the natural environment. A particularly important question is whether impacts initiated in early life might be exacerbated or attenuated over time in the presence or absence of other stressors in the field. We reared Olympia oyster (Ostrea lurida) larvae in laboratory cultures under control and elevated seawater pCO2 concentrations, quantified settlement success and size at metamorphosis, then outplanted juveniles to Tomales Bay, California, in the mid intertidal zone where emersion and temperature stress were higher, and in the low intertidal zone where conditions were more benign. We tracked survival and growth of outplanted juveniles for 4 months, halfway to reproductive age. Survival to metamorphosis in the laboratory was strongly affected by larval exposure to elevated pCO2 conditions. Survival of juvenile outplants was reduced dramatically at mid shore compared to low shore levels regardless of the pCO2 level that oysters experienced as larvae. However, juveniles that were exposed to elevated pCO2 as larvae grew less than control individuals, representing a larval carry-over effect. Although juveniles grew less at mid shore than low shore levels, there was no evidence of an interaction between the larval carry-over effect and shore level, suggesting little modulation of acidification impacts by emersion or temperature stress. Importantly, the carry-over effects of larval exposure to ocean acidification remained unabated 4 months later with no evidence of compensatory growth, even under benign conditions. This latter result points to the potential for extended consequences of brief exposures to altered seawater chemistry with potential consequences for population dynamics.

Persistent carry-over effects of planktonic exposure to ocean acidification in the Olympia oyster.

Predicting impacts of global environmental change is challenging due to the complex life cycles that characterize many terrestrial and aquatic taxa. Different life stages often interact with the physical environment in distinct ways, and a growing body of work suggests that stresses experienced during one life stage can "carry over" to influence subsequent stages. Assessments of population responses to environmental perturbation must therefore consider how effects might propagate across life-history transitions. We investigated consequences of ocean acidification (decreased pH and carbonate saturation) for early life stages of the Olympia oyster (Ostrea lurida), a foundation species in estuaries along the Pacific coast of North America. We reared oysters at three levels of seawater pH, including a control (8.0) and two additional levels (7.9 and 7.8). Oysters were cultured through their planktonic larval period to metamorphosis and into early juvenile life. Larvae reared under pH 7.8 exhibited a 15% decrease in larval shell growth rate, and a 7% decrease in shell area at settlement, compared to larvae reared under control conditions. Impacts were even more pronounced a week after settlement, with juveniles that had been reared as larvae under reduced pH exhibiting a 41% decrease in shell growth rate. Importantly, the latter effect arose regardless of the pH level the oysters experienced as juveniles, indicating a strong carry-over effect from the larval phase. Adverse impacts of early exposure to low pH persisted for at least 1.5 months after juveniles were transferred to a common environment. Overall, our results suggest that a stringent focus on a single phase of the life cycle (e.g., one perceived as the "weakest link") may neglect critical impacts that can be transferred across life stages in taxa with complex life histories.

Low-pH seawater alters indirect interactions in rocky-shore tidepools.

Ocean acidification is expected to degrade marine ecosystems, yet most studies focus on organismal-level impacts rather than ecological perturbations. Field studies are especially sparse, particularly ones examining shifts in direct and indirect consumer interactions. Here we address such connections within tidepool communities of rocky shores, focusing on a three-level food web involving the keystone sea star predator, Pisaster ochraceus, a common herbivorous snail, Tegula funebralis, and a macroalgal basal resource, Macrocystis pyrifera. We demonstrate that during nighttime low tides, experimentally manipulated declines in seawater pH suppress the anti-predator behavior of snails, bolstering their grazing, and diminishing the top-down influence of predators on basal resources. This attenuation of top-down control is absent in pools maintained experimentally at higher pH. These findings suggest that as ocean acidification proceeds, shifts of behaviorally mediated links in food webs could change how cascading effects of predators manifest within marine communities.

Functional impacts of ocean acidification in an ecologically critical foundation species.

Anthropogenic CO(2) is reducing the pH and altering the carbonate chemistry of seawater, with repercussions for marine organisms and ecosystems. Current research suggests that calcification will decrease in many species, but compelling evidence of impaired functional performance of calcium carbonate structures is sparse, particularly in key species. Here we demonstrate that ocean acidification markedly degrades the mechanical integrity of larval shells in the mussel Mytilus californianus, a critical community member on rocky shores throughout the northeastern Pacific. Larvae cultured in seawater containing CO(2) concentrations expected by the year 2100 (540 or 970 ppm) precipitated weaker, thinner and smaller shells than individuals raised under present-day seawater conditions (380 ppm), and also exhibited lower tissue mass. Under a scenario where mussel larvae exposed to different CO(2) levels develop at similar rates, these trends suggest a suite of potential consequences, including an exacerbated vulnerability of new settlers to crushing and drilling attacks by predators; poorer larval condition, causing increased energetic stress during metamorphosis; and greater risks from desiccation at low tide due to shifts in shell area to body mass ratios. Under an alternative scenario where responses derive exclusively from slowed development, with impacted individuals reaching identical milestones in shell strength and size by settlement, a lengthened larval phase could increase exposure to high planktonic mortality rates. In either case, because early life stages operate as population bottlenecks, driving general patterns of distribution and abundance, the ecological success of this vital species may be tied to how ocean acidification proceeds in coming decades.

Neuromuscular performance of paretic versus non-paretic plantar flexors after stroke.

The objective of this study was to compare the neuromuscular function of the paretic and non-paretic plantar flexors (i.e. soleus, gastrocnemius medialis, lateralis) in chronic stroke patients. It was hypothesized that the contractile rate of force development (RFD) and neural activation, assessed by electromyogram (EMG) and V-waves normalized to the M-wave, and voluntary activation (twitch interpolation) would be reduced during plantar flexor maximum voluntary isometric contraction and that the evoked muscle twitch properties would be reduced in the paretic limb. Ten chronic stroke survivors completed the study. The main findings were that the paretic side showed deteriorated function compared to the non-paretic leg in terms of (1) RFD in all analyzed time windows from force onset to 250 ms, although relative RFD (i.e. normalized to maximum voluntary force) was similar; (2) fast neural activation (for most analyzed time windows), assessed by EMG activity in time windows from EMG onset to 250 ms; (3) V-wave responses (except for gastrocnemius medialis); (4) voluntary activation; (5) the evoked peak twitch force, although there was no evidence of intrinsic muscle slowing; (6) EMG activity obtained at maximal voluntary force. In conclusion, this study demonstrates considerable neuromuscular asymmetry of the plantar flexors in chronic stroke survivors. Effective rehabilitation regimes should be investigated.