Seawater pH,and not inorganic nitrogen source,affects pH at the blade surface of Macrocystis pyrifera: implications for responses of the giant kelp to future oceanic conditions

Ocean acidification (OA), the ongoing decline in seawater pH, is predicted to have wide‐ranging effects on marine organisms and ecosystems. For seaweeds, the pH at the thallus surface, within the diffusion boundary layer (DBL), is one of the factors controlling their response to OA. Surface pH is controlled by both the pH of the bulk seawater and by the seaweeds' metabolism: photosynthesis and respiration increase and decrease pH within the DBL (pH_DBL), respectively. However, other metabolic processes, especially the uptake of inorganic nitrogen (N_i; NO₃^? and NH₄⁺) may also affect the pH_DBL. Using Macrocystis pyrifera, we hypothesized that (1) NO₃^? uptake will increase the pH_DBL, whereas NH₄⁺ uptake will decrease it, (2) if NO₃^? is cotransported with H⁺, increases in pH_DBL would be greater under an OA treatment (pH = 7.65) than under an ambient treatment (pH = 8.00), and (3) decreases in pH_DBL will be smaller at pH 7.65 than at pH 8.00, as higher external [H⁺] might affect the strength of the diffusion gradient. Overall, N_i source did not affect the pH_DBL. However, increases in pH_DBL were greater at pH 7.65 than at pH 8.00. CO₂ uptake was higher at pH 7.65 than at pH 8.00, whereas HCO₃^? uptake was unaffected by pH. Photosynthesis and respiration control pH_DBL rather than N_i uptake. We suggest that under future OA, Macrocystis pyrifera will metabolically modify its surface microenvironment such that the physiological processes of photosynthesis and N_i uptake will not be affected by a reduced pH.     

Bicarbonate uptake via an anion exchange protein is the main mechanism of inorganic carbon acquisition by the giant kelp Macrocystis pyrifera (Laminariales,Phaeophyceae) under variable pH

Macrocystis pyrifera is a widely distributed, highly productive, seaweed. It is known to use bicarbonate (HCO₃^?) from seawater in photosynthesis and the main mechanism of utilization is attributed to the external catalyzed dehydration of HCO₃^? by the surface‐bound enzyme carbonic anhydrase (CA_ext). Here, we examined other putative HCO₃^? uptake mechanisms in M. pyrifera under pH_T 9.00 (HCO₃^?: CO₂ = 940:1) and pH_T 7.65 (HCO₃^?: CO₂ = 51:1). Rates of photosynthesis, and internal CA (CA_int) and CA_ext activity were measured following the application of AZ which inhibits CA_ext, and DIDS which inhibits a different HCO₃^? uptake system, via an anion exchange (AE) protein. We found that the main mechanism of HCO₃^? uptake by M. pyrifera is via an AE protein, regardless of the HCO₃^?: CO₂ ratio, with CA_ext making little contribution. Inhibiting the AE protein led to a 55%–65% decrease in photosynthetic rates. Inhibiting both the AE protein and CA_ext at pH_T 9.00 led to 80%–100% inhibition of photosynthesis, whereas at pH_T 7.65, passive CO₂ diffusion supported 33% of photosynthesis. CA_int was active at pH_T 7.65 and 9.00, and activity was always higher than CA_ext, because of its role in dehydrating HCO₃^? to supply CO₂ to RuBisCO. Interestingly, the main mechanism of HCO₃^? uptake in M. pyrifera was different than that in other Laminariales studied (CA_ext‐catalyzed reaction) and we suggest that species‐specific knowledge of carbon uptake mechanisms is required in order to elucidate how seaweeds might respond to future changes in HCO₃^?:CO₂ due to ocean acidification.     

Assessing physiological tipping point of sea urchin larvae exposed to a broad range of pH

Our ability to project the impact of global change on marine ecosystem is limited by our poor understanding on how to predict species sensitivity. For example, the impact of ocean acidification is highly species‐specific, even in closely related taxa. The aim of this study was to test the hypothesis that the tolerance range of a given species to decreased pH corresponds to their natural range of exposure. Larvae of the green sea urchin Strongylocentrotus droebachiensis were cultured from fertilization to metamorphic competence (29 days) under a wide range of pH (from pH_T = 8.0/pCO₂ ≈ 480 μatm to pH_T = 6.5/pCO₂ ≈ 20 000 μatm) covering present (from pH_T 8.7 to 7.6), projected near‐future variability (from pH_T 8.3 to 7.2) and beyond. Decreasing pH impacted all tested parameters (mortality, symmetry, growth, morphometry and respiration). Development of normal, although showing morphological plasticity, swimming larvae was possible as low as pH_T ≥ 7.0. Within that range, decreasing pH increased mortality and asymmetry and decreased body length (BL) growth rate. Larvae raised at lowered pH and with similar BL had shorter arms and a wider body. Relative to a given BL, respiration rates and stomach volume both increased with decreasing pH suggesting changes in energy budget. At the lowest pHs (pH_T ≤ 6.5), all the tested parameters were strongly negatively affected and no larva survived past 13 days post fertilization. In conclusion, sea urchin larvae appeared to be highly plastic when exposed to decreased pH until a physiological tipping point at pH_T = 7.0. However, this plasticity was associated with direct (increased mortality) and indirect (decreased growth) consequences for fitness.     

The physiological response of marine diatoms to ocean acidification: differential roles of seawater pCO2 and pH

Although increasing the pCO₂ for diatoms will presumably down‐regulate the CO₂‐concentrating mechanism (CCM) to save energy for growth, different species have been reported to respond differently to ocean acidification (OA). To better understand their growth responses to OA, we acclimated the diatoms Thalassiosira pseudonana, Phaeodactylum tricornutum, and Chaetoceros muelleri to ambient (pCO₂ 400 μatm, pH 8.1), carbonated (pCO₂ 800 μatm, pH 8.1), acidified (pCO₂ 400 μatm, pH 7.8), and OA (pCO₂ 800 μatm, pH 7.8) conditions and investigated how seawater pCO₂ and pH affect their CCMs, photosynthesis, and respiration both individually and jointly. In all three diatoms, carbonation down‐regulated the CCMs, while acidification increased both the photosynthetic carbon fixation rate and the fraction of CO₂ as the inorganic carbon source. The positive OA effect on photosynthetic carbon fixation was more pronounced in C. muelleri, which had a relatively lower photosynthetic affinity for CO₂, than in either T. pseudonana or P. tricornutum. In response to OA, T. pseudonana increased respiration for active disposal of H⁺ to maintain its intracellular pH, whereas P. tricornutum and C. muelleri retained their respiration rate but lowered the intracellular pH to maintain the cross‐membrane electrochemical gradient for H⁺ efflux. As the net result of changes in photosynthesis and respiration, growth enhancement to OA of the three diatoms followed the order of C. muelleri > P. tricornutum > T. pseudonana. This study demonstrates that elucidating the separate and joint impacts of increased pCO₂ and decreased pH aids the mechanistic understanding of OA effects on diatoms in the future, acidified oceans.     

Seascape genetics of the stalked kelp Pterygophora californica and comparative population genetics in the Santa Barbara Channel

We conducted a population genetic analysis of the stalked kelp, Pterygophora californica, in the Santa Barbara Channel, California, USA. The results were compared with previous work on the genetic differentiation of giant kelp, Macrocystis pyrifera, in the same region. These two sympatric kelps not only share many life history and dispersal characteristics but also differ in that dislodged P. californica does not produce floating rafts with buoyant fertile sporophytes, commonly observed for M. pyrifera. We used a comparative population genetic approach with these two species to test the hypothesis that the ability to produce floating rafts increases the genetic connectivity among kelp patches in the Santa Barbara Channel. We quantified the association of habitat continuity and oceanographic distance with the genetic differentiation observed in stalked kelp, like previously conducted for giant kelp. We compared both overall (across all patches) and pairwise (between patches) genetic differentiation. We found that oceanographic transit time, habitat continuity, and geographic distance were all associated with genetic connectivity in P. californica, supporting similar previous findings for M. pyrifera. Controlling for differences in heterozygosity between kelp species using Jost's D_EST, we showed that global differentiation and pairwise differentiation were similar among patches between the two kelp species, indicating that they have similar dispersal capabilities despite their differences in rafting ability. These results suggest that rafting sporophytes do not play a significant role in effective dispersal of M. pyrifera at ecologically relevant spatial and temporal scales.     

Microsensor studies on Padina from a natural CO2 seep: implications of morphology on acclimation to low pH

Low seawater pH can be harmful to many calcifying marine organisms, but the calcifying macroalgae Padina spp. flourish at natural submarine carbon dioxide seeps where seawater pH is low. We show that the microenvironment created by the rolled thallus margin of Padina australis facilitates supersaturation of CaCO₃ and calcifi‐cation via photosynthesis‐induced elevated pH. Using microsensors to investigate oxygen and pH dynamics in the microenvironment of P. australis at a shallow CO₂ seep, we found that, under saturating light, the pH inside the microenvironment (pH_ME) was higher than the external seawater (pH_SW) at all pH_SW levels investigated, and the difference (i.e., pH_ME ? pH_SW) increased with decreasing pH_SW (0.9 units at pH_SW 7.0). Gross photosynthesis (P_g) inside the microenvironment increased with decreasing pH_SW, but algae from the control site reached a threshold at pH 6.5. Seep algae showed no pH threshold with respect to P_g within the pH_SW range investigated. The external carbonic anhydrase (CA) inhibitor, acetazolamide, strongly inhibited P_g of P. australis at pH_SW 8.2, but the effect was diminished under low pH_SW (6.4–7.5), suggesting a greater dependence on membrane‐bound CA for the dehydration of HCO₃^? ions during dissolved inorganic carbon uptake at the higher pH_SW. In comparison, a calcifying green alga, Halimeda cuneata f. digitata, was not inhibited by AZ, suggesting efficient bicarbonate transport. The ability of P. australis to elevate pH_ME at the site of calcification and its strong dependence on CA may explain why it can thrive at low pH_SW.     

Comparing salinity tolerance in early stages of the sporophytes of a non-indigenous kelp (Undaria pinnatifida) and a native kelp (Saccharina latissima)

The short-term effects of low salinities on the survival of germlings of an introduced kelp Undaria pinnatifida and a native kelp Saccharina latissima were assessed under laboratory conditions. This experiment was designed to compare the differential stress tolerance to salinity of the early life history stages of sporophytes of these two kelps that co-occur on European Atlantic coasts. Germlings (young sporophytes) of both species were exposed for 4 days to salinities ranging from 31 (control) to 26, 21, 16, 11, and 6 psu. Afterwards, they were post-cultured in control seawater (31 psu) for another 4 days to corroborate the viability of injured germlings. Results showed that germlings of the introduced kelp were less resistant to low salinity, surviving to as low as 16 psu; whereas the germlings of the native kelp survived in salinities as low as 11 psu. Despite the observed differences, both species are relatively tolerant to low salinity. Our observations also indicated that, at least in a short term, gametophytes of both species were able to survive in salinities as low as 6 psu. The significance of low-salinity tolerance to the distribution of these kelps and for their offshore cultivation is discussed.     

Sexual compatibility and hybrid formation between the giant kelp species <Emphasis Type="Italic">Macrocystis pyrifera</Emphasis> and <Emphasis Type="Italic">M. integrifolia</Emphasis> (Laminariales,Phaeophyceae) in Chile

Two species of giant kelp inhabit the coast of Chile: Macrocystis integrifolia and M. pyrifera, representing important economic resources. As part of our efforts to domesticate these kelps for mariculture, and to obtain superior cultivars, we studied their biological relationship. Hybridization experiments with clonal gametophyte cultures showed reciprocal cross-fertility and produced fertile hybrid sporophytes with intermediate morphological characters. This hybridization potential in the laboratory contrasts with the persistence of two morphologically well-defined sister taxa in natural habitats on the Pacific coast of South America. We conclude that M. integrifolia and M. pyrifera are conspecific and speculate that unknown mechanisms support the co-existence of two morphologically distinct taxa on the subspecific level.     

Bioturbation determines the response of benthic ammonia-oxidizing microorganisms to ocean acidification

Ocean acidification (OA), caused by the dissolution of increasing concentrations of atmospheric carbon dioxide (CO₂) in seawater, is projected to cause significant changes to marine ecology and biogeochemistry. Potential impacts on the microbially driven cycling of nitrogen are of particular concern. Specifically, under seawater pH levels approximating future OA scenarios, rates of ammonia oxidation (the rate-limiting first step of the nitrification pathway) have been shown to dramatically decrease in seawater, but not in underlying sediments. However, no prior study has considered the interactive effects of microbial ammonia oxidation and macrofaunal bioturbation activity, which can enhance nitrogen transformation rates. Using experimental mesocosms, we investigated the responses to OA of ammonia oxidizing microorganisms inhabiting surface sediments and sediments within burrow walls of the mud shrimp Upogebia deltaura. Seawater was acidified to one of four target pH values (pH_T 7.90, 7.70, 7.35 and 6.80) in comparison with a control (pH_T 8.10). At pH_T 8.10, ammonia oxidation rates in burrow wall sediments were, on average, fivefold greater than in surface sediments. However, at all acidified pH values (pH ≤ 7.90), ammonia oxidation rates in burrow sediments were significantly inhibited (by 79–97%; p < 0.01), whereas rates in surface sediments were unaffected. Both bacterial and archaeal abundances increased significantly as pH_T declined; by contrast, relative abundances of bacterial and archaeal ammonia oxidation (amoA) genes did not vary. This research suggests that OA could cause substantial reductions in total benthic ammonia oxidation rates in coastal bioturbated sediments, leading to corresponding changes in coupled nitrogen cycling between the benthic and pelagic realms.     

Sensitivity of sea urchin fertilization to pH varies across a natural pH mosaic

In the coastal ocean, temporal fluctuations in pH vary dramatically across biogeographic ranges. How such spatial differences in pH variability regimes might shape ocean acidification resistance in marine species remains unknown. We assessed the pH sensitivity of the sea urchin Strongylocentrotus purpuratus in the context of ocean pH variability. Using unique male–female pairs, originating from three sites with similar mean pH but different variability and frequency of low pH (pH_T ≤ 7.8) exposures, fertilization was tested across a range of pH (pH_T 7.61–8.03) and sperm concentrations. High fertilization success was maintained at low pH via a slight right shift in the fertilization function across sperm concentration. This pH effect differed by site. Urchins from the site with the narrowest pH variability regime exhibited the greatest pH sensitivity. At this site, mechanistic fertilization dynamics models support a decrease in sperm–egg interaction rate with decreasing pH. The site differences in pH sensitivity build upon recent evidence of local pH adaptation in S. purpuratus and highlight the need to incorporate environmental variability in the study of global change biology.     

Decreased photosynthesis and growth with reduced respiration in the model diatom Phaeodactylum tricornutum grown under elevated CO2 over 1800 generations

Studies on the long‐term responses of marine phytoplankton to ongoing ocean acidification (OA) are appearing rapidly in the literature. However, only a few of these have investigated diatoms, which is disproportionate to their contribution to global primary production. Here we show that a population of the model diatom Phaeodactylum tricornutum, after growing under elevated CO₂ (1000 μatm, HCL, pH_T: 7.70) for 1860 generations, showed significant differences in photosynthesis and growth from a population maintained in ambient CO₂ and then transferred to elevated CO₂ for 20 generations (HC). The HCL population had lower mitochondrial respiration, than did the control population maintained in ambient CO₂ (400 μatm, LCL, pH_T: 8.02) for 1860 generations. Although the cells had higher respiratory carbon loss within 20 generations under the elevated CO₂, being consistent to previous findings, they downregulated their respiration to sustain their growth in longer duration under the OA condition. Responses of phytoplankton to OA may depend on the timescale for which they are exposed due to fluctuations in physiological traits over time. This study provides the first evidence that populations of the model species, P. tricornutum, differ phenotypically from each other after having been grown for differing spans of time under OA conditions, suggesting that long‐term changes should be measured to understand responses of primary producers to OA, especially in waters with diatom‐dominated phytoplankton assemblages.     

Ocean acidification and seaweed reproduction: increased CO2 ameliorates the negative effect of lowered pH on meiospore germination in the giant kelp Macrocystis pyrifera (Laminariales,Phaeophyceae)

The worldwide effects of ocean acidification (OA) on marine species are a growing concern. In temperate coastal seas, seaweeds are dominant primary producers that create complex habitats and supply energy to higher trophic levels. Studies on OA and macroalgae have focused on calcifying species and adult stages, but critically, they have overlooked the microscopic stages of the reproductive life cycle, which, for other anthropogenic stressors, e.g., UV‐B radiation, are the most susceptible life‐history phase. Also, environmental cues and stressors can cause changes in the sex ratio, which has implications for the mating system and recruitment success. Here, we report the effects of pH (7.59–8.50) on meiospore germination and sex determination for the giant kelp, Macrocystis pyrifera (Laminariales), in the presence and absence of additional dissolved inorganic carbon (DIC). Lowered pH (7.59–7.60, using HCl‐only) caused a significant reduction in germination, whereas added DIC had the opposite effect, indicating that increased CO₂ at lower pH ameliorates physiological stress. This finding also highlights the importance of appropriate manipulation of seawater carbonate chemistry when testing the effects of OA on photosynthetic organisms. The proportion of male to female gametophytes did not vary significantly between treatments, suggesting that pH was not a primary environmental modulator of sex. Relative to the baseline (pH 8.19), gametophytes were 32% larger under moderate OA (pH 7.86) and 10% larger under extreme OA (pH 7.61). We suggest that metabolically active cells can compensate for the acidification of seawater. This homeostatic function minimizes the negative effects of lower pH (high H⁺ ions) on cellular activity. The 6–9% reduction in germination success under extreme OA suggests that meiospores of M. pyrifera may be resistant to future OA.     

Seasonal variation in epifaunal communities associated with giant kelp (Macrocystis pyrifera) at an upwelling‐dominated site

Kelp forests are highly productive and species‐rich benthic ecosystems in temperate regions that provide biogenic habitat for numerous associated species. Diverse epifaunal communities inhabit kelp sporophytes and are subject to variations in the physical environment and to changes experienced by the kelp habitat itself. We assessed seasonal variations in epifaunal invertebrate communities inhabiting giant kelps, Macrocystis pyrifera, and their effects on this seaweed. Six seasonal samplings were conducted over a year at an upwelling‐dominated site in northern‐central Chile where physical conditions are known to fluctuate temporally. More than 30 taxa were identified, among which peracarid crustaceans stood out in both diversity and abundance. Species richness and abundance differed among sporophyte sections (holdfast and fronds) and throughout the year. The frond community was dominated by two grazers (the amphipod Peramphithoe femorata and the isopod Amphoroidea typa), while suspension feeders, grazers, and omnivores (the amphipod Aora typica, the isopod Limnoria quadripunctata, and polychaetes) dominated the holdfasts. Abundances of the dominant species fluctuated throughout the year but patterns of variation differed among species. The most abundant grazer (P. femorata) had highest densities in summer, while the less abundant grazer (A. typa) reached its peak densities in winter. Interestingly, the area of kelp damaged by grazers was highest in autumn and early winter, suggesting that grazing impacts accumulate during periods of low kelp growth, which can thus be considered as ‘vestiges of herbivory past.’ Among the factors determining the observed seasonal patterns, strong variability of environmental conditions, reproductive cycles of associated fauna, and predation by fishes vary in importance. Our results suggest that during spring and early summer, bottom‐up processes shape the community structure of organisms inhabiting large perennial seaweeds, whereas during late summer and autumn, top‐down processes are more important.     

THE EFFECTS OF HIGH IRRADIANCE ON THE SETTLEMENT COMPETENCY AND VIABILITY OF KELP ZOOSPORES1

Elevated irradiance has a profound effect on the successful dispersal and establishment of kelp zoospores, affecting their physiology and viability. The research to date, however, has been on zoospores localized near the benthos, with little attention on the importance of vertical transportation and subsequent exposure to increased irradiance. Therefore, we wanted to investigate the effects of exposure to high irradiance on the reproductive planktonic life‐history stages of kelps Macrocystis pyrifera (L.) C. Agardh and Pterygophora californica Rupr. Zoospores of both species were exposed to different irradiances (75, 275, 575, 1,025 μmol photons · m^?2 · s^?1) over varying durations (1, 2, 4, 8, 12 h) and subsequently monitored for settlement competency, gametophyte development, and reproductive viability. Settlement success for M. pyrifera was uniform throughout all irradiance × time treatments, while settlement for P. californica decreased with increasing exposure time but not irradiance, although settlement was generally reduced at the highest irradiance level. Following zoospore settlement, germ tube development was visible in the gametophytes of both species within 1 week, although a significant decline of germ tube density in P. californica was observed with increasing irradiance. Similarly, a decrease in germ tube development with increasing exposure was observed across all irradiance levels for M. pyrifera, but irradiance itself was not significant. Further development into embryonic sporophytes was remarkably similar to gametophyte development, suggesting that the effect of exposure of kelp zoospores to high irradiance on subsequent sporophyte production is mediated through gametophyte development as well as zoospore survival.     

Biosorption of chromium(III) by two brown algae Macrocystis pyrifera and Undaria pinnatifida: Equilibrium and kinetic study

Two brown algae, Macrocystis pyrifera and Undaria pinnatifida, were employed to remove Cr(III) from aqueous solutions. Both seaweeds were characterized in terms of alginate yields. The alginate contents were 20 and 30% of the dry weight for M. pyrifera and U. pinnatifida, respectively. Kinetics experiments were carried out at different initial pH values. Cr(III) biosorption was affected by the solution pH. The highest metal uptake was found at pH 4 for both biosorbents. Different models were applied to elucidate the rate‐controlling mechanism: pseudo‐first‐order, pseudo‐second‐order, external mass transfer and intra‐particle diffusion. The application of Langmuir, Freundlich and Dubinin–Radushkevich models to the equilibrium data showed a better fitting to the first model. The maximum Cr(III) sorption capacity (q_m) and the affinity coefficient (b) were very similar for both biosorbents: 0.77 mmol/g and 1.20 L/mmol for M. pyrifera and 0.74 mmol/g and 1.06 L/mmol for U. pinnatifida. The free energy of the sorption process was estimated using the Dubinin–Radushkevich isotherm. The values indicate that the processes are chemical sorptions. To evaluate the significance of the ion‐exchange mechanism, the light metals (Ca²⁺, Na⁺, Mg²⁺ and K⁺) and pH were measured during the experiments.     

Compensatory abilities depending on seasonal timing of thallus excision of the kelp Undaria pinnatifida cultivated in Matsushima Bay, northern Japan

Improved cultivation technology for the kelp Undaria pinnatifida is greatly needed to increase production to meet increasing commercial demand. A previous cultivation trial indicated that the crop yield of U. pinnatifida sporophytes could be increased greatly by thallus excision in late February due to compensatory growth of the remaining tissues. To develop this potential new cultivation technology, it is essential to identify the time period during which this kelp can compensate and its physiological responses to thallus excision. In this study, U. pinnatifida sporophytes were excised at about 30 cm above the meristem at the beginning of January, February, March, and April, respectively. Morphological features, photosynthesis, nutrient uptake, and carbon and nitrogen contents of excised kelps were measured and compared with these parameters in control kelps grown without excision. Both experimental and control kelps were farmed together in Matsushima Bay, northern Japan. The kelps excised in January and February showed significant increases in the lengths and dry weights of the blade, photosynthetic rates, nutrient uptake rates, and carbon and nitrogen contents compared with the control kelps, and the growth phase was prolonged for at least 1 month. No significant increases were found in dry weights and carbon and nitrogen contents of sporophylls until early April, which indicated that the maturation period was delayed. At the end of this experiment, the nitrogen contents of sporophyll tissues formed after excisions were significantly lower than those of tissues formed before excisions. In contrast, the kelps excised in March and April showed no significant increases in morphological and physiological parameters compared with control kelps. These results suggest that U. pinnatifida sporophytes exhibited great compensation when excisions were conducted during the growing phase in January and February but not in March and April when the maturation phase had started. The regulation of resource allocation to growth and maturation after thallus excisions in January and February likely results in prolongation of the growth phase and maturation phase in excised kelps.     

Effects of increased pCO2 on zinc uptake and calcification in the tropical coral Stylophora pistillata

Zinc (Zn) is an essential element for corals. We investigated the effects of ocean acidification on zinc incorporation, photosynthesis, and gross calcification in the scleractinian coral Stylophora pistillata. Colonies were maintained at normal pH_T (8.1) and at two low-pH conditions (7.8 and 7.5) for 5 weeks. Corals were exposed to ⁶⁵Zn dissolved in seawater to assess uptake rates. After 5 weeks, corals raised at pH_T (8.1) exhibited higher ⁶⁵Zn activity in the coral tissue and skeleton, compared with corals raised at a lower pH. Photosynthesis, photosynthetic efficiency, and gross calcification, measured by ⁴⁵Ca incorporation, were however unchanged even at the lowest pH.     

Development of PCR‐Based Markers to Determine the Sex of Kelps

Sex discriminating genetic markers are commonly used to facilitate breeding programs in economically and ecologically important animal and plant species. However, despite their considerable economic and ecological value, the development of sex markers for kelp species has been very limited. In this study, we used the recently described sequence of the sex determining region (SDR) of the brown algal model Ectocarpus to develop novel DNA-based sex-markers for three commercially relevant kelps: Laminaria digitata, Undaria pinnatifida and Macrocystis pyrifera. Markers were designed within nine protein coding genes of Ectocarpus male and female (U/V) sex chromosomes and tested on gametophytes of the three kelp species. Seven primer pairs corresponding to three loci in the Ectocarpus SDR amplified sex-specific bands in the three kelp species, yielding at least one male and one female marker for each species. Our work has generated the first male sex-specific markers for L. digitata and U. pinnatifida, as well as the first sex markers developed for the genus Macrocystis. The markers and methodology presented here will not only facilitate seaweed breeding programs but also represent useful tools for population and demography studies and provide a means to investigate the evolution of sex determination across this largely understudied eukaryotic group.     

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 (Ω_ara) 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, Ω_ara 1.29), and nor are the rates of skeletal dissolution in D. dianthus. We transplanted D. dianthus to 350 m depth (pH_T 8.02; pCO₂ 448 μatm, Ω_ara 2.58) and to a 3 m depth CO₂ seep in oligotrophic waters (pH_T 7.35; pCO₂ 2879 μatm, Ω_ara 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.     

In a squeeze: Epibiosis may affect the distribution of kelp forests

The processes limiting the population recovery of the kelp Saccharina latissima after recent large‐scale loss from the south coast of Norway are poorly understood. Previous investigations do, however, suggest that the impacts of biotic interactions (epibiosis and competition) and increased water turbidity are important. We investigated the depth‐related patterns of growth, epibiosis, and mortality in two sample populations of kelp, from the south and the southwest coast of Norway. The investigations were performed over a period of seven months, in a crossed translocational study, where kelps were mounted on rigs at six depths (1, 3, 6, 9, 15, and 24 m). In a second experiment, the amounts of light blocked by different epibiont layers growing on the kelp frond were investigated. While growth decreased with depth in spring and summer, the kelp grew faster at 15 m than at shallower depths in fall. Survival was low both in shallow water and below 15 m depth. Epibionts covered the kelp growing at depths from 1 to 9 m, and the laboratory study showed that the coverage may have deprived the individuals of as much as 90% of the available light. Although the depth‐related results we present apply—in the strictest sense—only to kelp translocated on rigs, we argue that the relative patterns are relevant for natural populations. Growth and survival of S. latissima is likely to be reduced by heavy loads of epibionts, while depths where epibionts are sparse may be close to the lower limit of the kelps depth distribution along the south coast of Norway. This suggests that a vertical squeeze, or narrowing of the distribution range of kelp forests may be occurring in Norway.