Scale-dependent topographic complexity underpins abundance and spatial distribution of ecosystem engineers on natural and artificial structures.

In response to ongoing coastal urbanization, it is critical to develop effective methods to improve the biodiversity and ecological sustainability of artificial shorelines. Enhancing the topographic complexity of coastal infrastructure through the mimicry of natural substrata may facilitate the establishment of ecosystem engineering species and associated biogenic habitat formation. However, interactions between ecosystem engineers and their substratum are likely determined by organismal size and resource needs, thus making responses to topography highly scale-dependent. Here, we assessed the topographic properties (rugosity, surface area, micro-surface orientations) that underpin the abundance and distribution of two ecosystem engineers (fucoids, limpets) across six spatial scales (1-500 mm). Furthermore, we assessed the 'biogenic' rugosity created by barnacle matrices across fine scales (1-20 mm). Field surveys and 3D scanning, conducted across natural and artificial substrata, showed major effects of rugosity and associated topographic variables on ecosystem engineer assemblages and spatial occupancy, while additional abiotic environmental factors (compass direction, wave exposure) and biotic associations only had weak influences. Natural substrata exhibited ≤67 % higher rugosity than artificial ones. Fucoid-covered patches were predominantly associated with high-rugosity substrata and horizontal micro-surfaces, while homescars of limpets (≥15 mm shell length) predominated on smoother substratum patches. Barnacle-driven rugosity homogenized substrata at scales ≤10 mm. Our findings suggest that scale-dependent rugosity is a key driver of fucoid habitat formation and limpet habitat use, with wider eco-engineering applications for mimicking ecologically impactful topography on coastal infrastructure.

Topography-based modulation of environmental factors as a mechanism for intertidal microhabitat formation: A basis for marine ecological design.

Topographic complexity is often considered to be closely associated with habitat complexity and niche diversity; however, complex topography per se does not imply habitat suitability. Rather, ecologically suitable habitats may emerge if topographic features interact with environmental factors and thereby alter their surrounding microenvironment to the benefit of local organisms (e.g., resource provisioning, stress mitigation). Topography may thus act as a key modulator of abiotic stressors and biotic pressures, particularly in environmentally challenging intertidal systems. Here, we review how topography can alter microhabitat conditions with respect to four resources required by intertidal organisms: a source of energy (light, suspended food particles, prey, detritus), water (hydration, buffering of light, temperature and hydrodynamics), shelter (temperature, wave exposure, predation), and habitat space (substratum area, propagule settlement, movement). We synthesize mechanisms and quantitative findings of how environmental factors can be altered through topography and suggest an organism-centered 'form-follows-ecological-function' approach to designing multifunctional marine infrastructure.

The north-south divide? Macroalgal functional trait diversity and redundancy varies with intertidal aspect.

BACKGROUND AND AIMS: Marine macroalgae ('seaweeds') are critical to coastal ecosystem structure and function, but also vulnerable to the many environmental changes associated with anthropogenic climate change (ACC). The local habitat conditions underpinning observed and predicted ACC-driven changes in intertidal macroalgal communities are complex and probably site-specific and operate in addition to more commonly reported regional factors such as sea surface temperatures. METHODS: We examined how the composition and functional trait expression of macroalgal communities in SW England varied with aspect (i.e. north-south orientation) at four sites with opposing Equator- (EF) and Pole-facing (PF) surfaces. Previous work at these sites had established that average annual (low tide) temperatures vary by 1.6 °C and that EF-surfaces experience six-fold more frequent extremes (i.e. >30 °C). KEY RESULTS: PF macroalgal communities were consistently more taxon rich; 11 taxa were unique to PF habitats, with only one restricted to EF. Likewise, functional richness and dispersion were greater on PF-surfaces (dominated by algae with traits linked to rapid resource capture and utilization, but low desiccation tolerance), although differences in both taxon and functional richness were probably driven by the fact that less diverse EF-surfaces were dominated by desiccation-tolerant fucoids. CONCLUSIONS: Although we cannot disentangle the influence of temperature variation on algal ecophysiology from the indirect effects of aspect on species interactions (niche pre-emption, competition, grazing, etc.), our study system provides an excellent model for understanding how environmental variation at local scales affects community composition and functioning. By virtue of enhanced taxonomic diversity, PF-aspects supported higher functional diversity and, consequently, greater effective functional redundancy. These differences may imbue PF-aspects with resilience against environmental perturbation, but if predicted increases in global temperatures are realized, some PF-sites may shift to a depauperate, desiccation-tolerant seaweed community with a concomitant loss of functional diversity and redundancy.

Invasive macroalgae in native seagrass beds: vectors of spread and impacts.

BACKGROUND AND AIMS: Worldwide, invasive species are spreading through marine systems at an unprecedented rate with both positive and negative consequences for ecosystems and the biological functioning of organisms. Human activities from shipping to habitat damage and modification are known vectors of spread, although biological interactions including epibiosis are increasingly recognized as potentially important to introduction into susceptible habitats. METHODS: We assessed a novel mechanism of spread - limpets as transporters of an invasive alga, Sargassum muticum, into beds of the seagrass Zostera marina - and the physiological impact of its invasion. The association of S. muticum with three limpet species and other habitats was assessed using intertidal surveys on rocky shores and snorkelling at two seagrass sites in the UK. A 4-year field study tested the effect of S. muticum on Z. marina shoot density, dry weight and phenolic compounds (caffeic and tannic acid) content, and a laboratory experiment tested the impact of S. muticum on nutrient partitioning (C/H/N/P/Si), photosynthetic efficiency (Fv/Fm) and growth of Z. marina. RESULTS: On rocky shores 15 % of S. muticum occurrences were attached to the shells of live limpets. In seagrass beds 5 % of S. muticum occurrences were attached to the shells of dead limpets. The remainder were attached to rock, to cobblestones, to the seagrass matrix or embedded within the sand. Z. marina density and phenolics content was lower when S. muticum co-occurred with it. Over 3 years, photosynthetic responses of Z. marina to S. muticum were idiosyncratic, and S. muticum had no effect on nutrient partitioning in Z. marina. CONCLUSIONS: Our results show limpets support S. muticum as an epibiont and may act as a previously unreported transport mechanism introducing invaders into sensitive habitats. S. muticum reduced production of phenolics in Z. marina, which may weaken its defensive capabilities and facilitate proliferation of S. muticum. The effect of S. muticum on Z. marina photosynthesis requires further work but having no effect on the capacity of Z. marina to sequester nutrients suggests a degree of resilience to this invader.

Victim of changes? Marine macroalgae in a changing world.

BACKGROUND: Marine macroalgae ('seaweeds') are a diverse and globally distributed group of photosynthetic organisms that together generate considerable primary productivity, provide an array of different habitats for other organisms, and contribute many important ecosystem functions and services. As a result of continued anthropogenic stress on marine systems, many macroalgal species and habitats face an uncertain future, risking their vital contribution to global productivity and ecosystem service provision. SCOPE: After briefly considering the remarkable taxonomy and ecological distribution of marine macroalgae, we review how the threats posed by a combination of anthropogenically induced stressors affect seaweed species and communities. From there we highlight five critical avenues for further research to explore (long-term monitoring, use of functional traits, focus on early ontogeny, biotic interactions and impact of marine litter on coastal vegetation). CONCLUSIONS: Although there are considerable parallels with terrestrial vascular plant responses to the many threats posed by anthropogenic stressors, we note that the impacts of some (e.g. habitat loss) are much less keenly felt in the oceans than on land. Nevertheless, and in common with terrestrial plant communities, the impact of climate change will inevitably be the most pernicious threat to the future persistence of seaweed species, communities and service provision. While understanding macroalgal responses to simultaneous environmental stressors is inevitably a complex exercise, our attempt to highlight synergies with terrestrial systems, and provide five future research priorities to elucidate some of the important trends and mechanisms of response, may yet offer some small contribution to this goal.

Marine heatwave duration and intensity interact to reduce physiological tipping points of kelp species with contrasting thermal affinities.

BACKGROUND AND AIMS: Marine heatwaves (MHWs) are widely recognized as pervasive drivers of ecosystem change, yet our understanding of how different MHW properties mediate ecological responses remains largely unexplored. Understanding MHW impacts on foundation species is particularly important, given their structural role in communities and ecosystems. METHODS: We simulated a series of realistic MHWs with different levels of intensity (Control: 14 °C, Moderate: 18 °C, Extreme: 22 °C) and duration (14 or 28 d) and examined responses of two habitat-forming kelp species in the southwest UK. Here, Laminaria digitata reaches its trailing edge and is undergoing a range contraction, whereas Laminaria ochroleuca reaches its leading edge and is undergoing a range expansion. KEY RESULTS: For both species, sub-lethal stress responses induced by moderate-intensity MHWs were exacerbated by longer duration. Extreme-intensity MHWs caused dramatic declines in growth and photosynthetic performance, and elevated bleaching, which were again exacerbated by longer MHW duration. Stress responses were most pronounced in L. ochroleuca, where almost complete tissue necrosis was observed by the end of the long-duration MHW. This was unexpected given the greater thermal safety margins assumed with leading edge populations. It is likely that prolonged exposure to sub-lethal thermal stress exceeded a physiological tipping point for L. ochroleuca, presumably due to depletion of internal reserves. CONCLUSIONS: Overall, our study showed that exposure to MHW profiles projected to occur in the region in the coming decades can have significant deleterious effects on foundation kelp species, regardless of their thermal affinities and location within respective latitudinal ranges, which would probably have consequences for entire communities and ecosystems.

Evaluation of Simvastatin as a Disease-Modifying Treatment for Patients With Parkinson Disease: A Randomized Clinical Trial.

Importance: Current treatments manage symptoms of Parkinson disease (PD), but no known treatment slows disease progression. Preclinical and epidemiological studies support the potential use of statins as disease-modifying therapy. Objective: To determine whether simvastatin has potential as a disease-modifying treatment for patients with moderate PD. Design, Setting, and Participants: This randomized clinical trial, a double-blind, parallel-group, placebo-controlled futility trial, was conducted between March 2016 and May 2020 within 23 National Health Service Trusts in England. Participants aged 40 to 90 years with a diagnosis of idiopathic PD, with a modified Hoehn and Yahr stage of 3.0 or less while taking medication, and taking dopaminergic medication with wearing-off phenomenon were included. Data were analyzed from May 2020 to September 2020, with additional analysis in February 2021. Interventions: Participants were allocated 1:1 to simvastatin or matched placebo via a computer-generated random sequence, stratified by site and Hoehn and Yahr stage. In the simvastatin arm, participants entered a 1-month phase of simvastatin, 40 mg daily, followed by 23 months of simvastatin, 80 mg daily, before a 2-month washout period. Main Outcomes and Measures: The prespecified primary outcome was 24-month change in Movement Disorder Society Unified Parkinson Disease Rating Scale (MDS-UPDRS) part III score measured while not taking medication (high scores indicate worse outcome). The primary futility analysis included participants who commenced the 80-mg phase and had valid primary outcome data. The safety analysis included all participants who commenced trial treatment and is reported by dose at time of event. Results: Of 332 patients assessed for eligibility, 32 declined and 65 were ineligible. Of 235 recruited participants, 97 (41%) were female, 233 (99%) were White, and the mean (SD) age was 65.4 (9.4) years. A total of 216 patients progressed to the 80-mg dose. Primary outcome analysis (n = 178) indicated the simvastatin group had an additional deterioration in MDS-UPDRS III score while not taking medication at 24 months compared with the placebo group (1.52 points; 2-sided 80% CI, -0.77 to 3.80; 1-sided futility test P = .006). A total of 37 serious adverse events (AEs), including 3 deaths, and 171 AEs were reported for participants receiving 0-mg simvastatin; 37 serious AEs and 150 AEs were reported for participants taking 40 mg or 80 mg of simvastatin. Four participants withdrew from the trial because of an AE. Conclusions and Relevance: In this randomized clinical trial, simvastatin was futile as a disease-modifying therapy in patients with PD of moderate severity, providing no evidence to support proceeding to a phase 3 trial. Trial Registration: ISRCTN Identifier: 16108482.

Climate-driven shifts in kelp forest composition reduce carbon sequestration potential.

The potential contribution of kelp forests to blue carbon sinks is currently of great interest but interspecific variance has received no attention. In the temperate Northeast Atlantic, kelp forest composition is changing due to climate-driven poleward range shifts of cold temperate Laminaria digitata and Laminaria hyperborea and warm temperate Laminaria ochroleuca. To understand how this might affect the carbon sequestration potential (CSP) of this ecosystem, we quantified interspecific differences in carbon export and decomposition alongside changes in detrital photosynthesis and biochemistry. We found that while warm temperate kelp exports up to 71% more carbon per plant, it decomposes up to 155% faster than its boreal congeners. Elemental stoichiometry and polyphenolic content cannot fully explain faster carbon turnover, which may be attributable to contrasting tissue toughness or unknown biochemical and structural defenses. Faster decomposition causes the detrital photosynthetic apparatus of L. ochroleuca to be overwhelmed 20 days after export and lose integrity after 36 days, while detritus of cold temperate species maintains carbon assimilation. Depending on the photoenvironment, detrital photosynthesis could further exacerbate interspecific differences in decomposition via a potential positive feedback loop. Through compositional change such as the predicted prevalence of L. ochroleuca, ocean warming may therefore reduce the CSP of such temperate marine forests.

On the diversity and distribution of a data deficient habitat in a poorly mapped region: The case of Sabellaria alveolata L. in Ireland.

Data that can be used to monitor biodiversity through time are essential for conservation and management. The reef-forming worm, Sabellaria alveolata (L. 1767) is currently classed as 'Data Deficient' due to an imbalance in the spread of data on its distribution. Little is known about the distribution of this species around Ireland. Using data archaeology, we collated past and present distribution records and discovered that S. alveolata has a discontinuous distribution with large gaps between populations. Many regions lack data and should be targeted for sampling. Biodiversity surveys revealed that S. alveolata supported diverse epibiotic algal communities. Retrograding (declining) reefs supported greater infaunal diversity than prograding (growing) reefs or sand, suggesting that S. alveolata is a dynamic ecosystem engineer that has a lasting legacy effect. Similar research should be carried out for other Data Deficient species, habitats and regions. Such data are invaluable resources for management and conservation.

Sustained productivity and respiration of degrading kelp detritus in the shallow benthos: Detached or broken, but not dead.

Temperate kelp forests contribute significantly to marine primary productivity and fuel many benthic and pelagic food chains. A large proportion of biomass is exported from kelp forests as detritus into recipient marine ecosystems, potentially contributing to Blue Carbon sequestration. The degradation of this organic material is slow and recent research has revealed the preservation of photosynthetic functions over time. However, the physiological correlates of detrital breakdown in Laminaria spp. have not yet been studied. The warming climate threatens to reshuffle the species composition of kelp forests and perturb the dynamics of these highly productive ecosystems. The present study compares the physiological response of degrading detritus from two competing North East Atlantic species; the native Boreal Laminaria hyperborea and the thermally tolerant Boreal-Lusitanian L. ochroleuca. Detrital fragment degradation was measured by a mesocosm experiment across a gradient of spectral attenuation (a proxy for depth) to investigate the changes in physiological performance under different environmental conditions. Degradation of fragments was quantified over 108 days by measuring the biomass, production and respiration (by respirometry) and efficiency of Photosystem II (by PAM fluorometry). Data indicated that whilst degrading, the photosynthetic performance of the species responded differently to simulated depths, but fragments of both species continued to produce oxygen for up to 56 days and sustained positive net primary production. This study reveals the potential for ostensibly detrital kelp to contribute to Blue Carbon fixation through sustained primary production which should be factored into Blue Carbon management. Furthermore, the physiological response of kelp detritus is likely dependent upon the range of habitats to which it is exported. In the context of climate change, shifts in species composition of kelp forests and their detritus are likely to have wide-reaching effects upon the cycling of organic matter in benthic ecosystems.

Simvastatin as a neuroprotective treatment for Parkinson's disease (PD STAT): protocol for a double-blind, randomised, placebo-controlled futility study.

INTRODUCTION: Parkinson's disease (PD) is a progressive neurodegenerative condition affecting approximately 185,000 people in the UK. No drug has been proven to slow disease progression. Epidemiological and pre-clinical data support simvastatin, a widely used cholesterol-lowering drug with a well-established safety profile, having neuroprotective properties. The aim of this study (Simvastatin as a neuroprotective treatment for PD (PD STAT)) is to determine whether simvastatin has the potential to slow PD progression. The study is part of the International Linked Clinical Trials initiative coordinated by The Cure Parkinson's Trust. This paper describes the protocol for the PD STAT study. METHODS AND ANALYSIS: PD STAT is a double-blind, randomised, placebo-controlled, multi-centre, parallel group, futility trial in patients with PD of mild-moderate severity. 235 participants have been recruited and randomly allocated in a 1:1 ratio to receive either oral simvastatin or matched placebo. Treatment involves a 1-month low-dose phase (40 mg daily), followed by a 23-month high-dose phase (80 mg daily) and ends with a 2-month washout period. Participants are reviewed at clinic visits at 1 month, 6, 12, 18, 24 and 26 months post-baseline, with interim telephone follow-up to monitor for adverse events.The primary outcome is the change in the Movement Disorder Society Unified Parkinson's Disease Rating Scale part III motor subscale score in the practically defined OFF medication state (OFF state) between baseline and 24 months. Primary analysis will be on a modified intention to treat basis and will include only those participants who progress to the high-dose phase of the study. ETHICS AND DISSEMINATION: The protocol has been approved by the North East-Newcastle and North Tyneside 2 Research Ethics Committee. The results will be disseminated via research articles in peer-reviewed journals and presentations at local, national and international scientific meetings, as well as disseminated via patient groups, websites and networks. A summary of the study findings will be posted to participants at the end of the study. TRIAL REGISTRATION: ISRCTN16108482 (prospectively registered); EudraCT 2015-000148-40; ClinicalTrials.gov NCT02787590; Pre-results.

Characterising the deterioration of different plastics in air and seawater.

In situ studies of plastic deterioration can help us understand the longevity of macroplastic as well as the generation of microplastics in the environment. Photo-oxidation contributing to the generation of microplastics in the marine environment was explored using four types of plastic (polyethene, polystyrene, poly(ethylene terephthalate) and Biothene® exposed in light and in shade, in both air and sea water. Metrics for deterioration were tensile extensibility and oxidation rate. Measurements were conducted at intervals between 7 and 600 days' exposure. Deterioration was faster in air than in sea water and was further accelerated in direct light compared to shade. Extensibility and oxidation were significantly inversely correlated in samples exposed in air. Samples in sea water lost extensibility at a slower rate. Polystyrene, which enters the waste stream rapidly due to its wide application in packaging, deteriorated fastest and is, therefore, likely to form microplastics more rapidly than other materials, especially when exposed to high levels of irradiation, for example when stranded on the shore.

Removal of intertidal grazers by human harvesting leads to alteration of species interactions, community structure and resilience to climate change.

Extreme fluctuations in abiotic conditions can induce a biological stress response (e.g. bleaching) detrimental to an organism's health. In some instances, organisms can recover if conditions are alleviated, such as through co-occurrence with other species that confer protection. Biodiverse, multitrophic communities are increasingly recognised as important promoters of species persistence and resilience under environmental change. On intertidal shores, the role of grazers as top-down determinants of algal community structure is well recognised. Similarly, the harvesting of grazers for human consumption is increasingly prevalent with potential to greatly alter the community dynamics. Here, we assess how differences in harvesting pressure of grazers under three management regimes (no-take; managed access; open-access) alters the trophic interactions between grazers, and algal communities. Grazer density and body size frequencies were different among regimes leading to changes in the photosynthetic performance and recovery of crustose coralline algae (CCA) post-bleaching, as well as their presence altering the strength of interactions between species. The exclusion of grazers from patches using cages led to different emergent communities and reduced negative correlations between taxa. The absence of larger grazers (>9 cm) at the managed access site led to macroalgal overgrowth of bleached CCA negatively affecting its recovery, whereas no-take or open-access led to a moderated algal growth and a shift from competitive to facilitative interactions between algal species. Given that CCA play an important role in the population growth and development of other species, the choice of management measure should be carefully considered before implementation, depending on objectives.

Individual and population-level responses to ocean acidification.

Ocean acidification is predicted to have detrimental effects on many marine organisms and ecological processes. Despite growing evidence for direct impacts on specific species, few studies have simultaneously considered the effects of ocean acidification on individuals (e.g. consequences for energy budgets and resource partitioning) and population level demographic processes. Here we show that ocean acidification increases energetic demands on gastropods resulting in altered energy allocation, i.e. reduced shell size but increased body mass. When scaled up to the population level, long-term exposure to ocean acidification altered population demography, with evidence of a reduction in the proportion of females in the population and genetic signatures of increased variance in reproductive success among individuals. Such increased variance enhances levels of short-term genetic drift which is predicted to inhibit adaptation. Our study indicates that even against a background of high gene flow, ocean acidification is driving individual- and population-level changes that will impact eco-evolutionary trajectories.

Calcification is not the Achilles' heel of cold-water corals in an acidifying ocean.

Ocean acidification is thought to be a major threat to coral reefs: laboratory evidence and CO2 seep research has shown adverse effects on many coral species, although a few are resilient. There are concerns that cold-water corals are even more vulnerable as they live in areas where aragonite saturation (Ωara ) is lower than in the tropics and is falling rapidly due to CO2 emissions. Here, we provide laboratory evidence that net (gross calcification minus dissolution) and gross calcification rates of three common cold-water corals, Caryophyllia smithii, Dendrophyllia cornigera, and Desmophyllum dianthus, are not affected by pCO2 levels expected for 2100 (pCO2  1058 µatm, Ωara 1.29), and nor are the rates of skeletal dissolution in D. dianthus. We transplanted D. dianthus to 350 m depth (pHT 8.02; pCO2  448 µatm, Ωara 2.58) and to a 3 m depth CO2 seep in oligotrophic waters (pHT 7.35; pCO2  2879 µatm, Ωara 0.76) and found that the transplants calcified at the same rates regardless of the pCO2 confirming their resilience to acidification, but at significantly lower rates than corals that were fed in aquaria. Our combination of field and laboratory evidence suggests that ocean acidification will not disrupt cold-water coral calcification although falling aragonite levels may affect other organismal physiological and/or reef community processes.