Brezo Martínez

SEASONAL VARIATION OF P CONTENT AND MAJOR N POOLS IN PALMARIA PALMATA (RHODOPHYTA)1

The annual variation of major nitrogen pools, phosphorus, carbon, ash, and thallus water content in relation to seasonal environmental changes was studied in two northern Spanish populations of the edible seaweed Palmaria palmata (Linnaeus) Kuntze. Observed patterns were investigated using Spearman rank order correlation coefficients. There were significant relationships between thallus nutrient content and nitrate and orthophosphate seawater concentration, irradiance, temperature, and wave force. The highest levels of total N and P and nitrogenous compounds were observed during autumn and winter because the thallus stored N‐ and P‐rich compounds in response to high nutrient seawater concentration when growth was limited by low light and temperature. Phycoerythrin and other proteins were the main N reserves. Thallus P content was higher in algae from the eutrophic site. During spring, reduced N and P thallus content and increased ash, water, and C content were observed in the growing fronds. N and P seawater concentrations were undetectable during summer when nutrient reserves were low and growth was reduced and eventually suppressed, suggesting nutrient limiting conditions. Palmaria palmata clearly could take advantage of elevated N and P concentrations to create storage reserves in winter to support early summer growth. This storage response reduced the dependence of algal nutrition on the external nutrient supply and supports the use of pulse fertilization to diminish summer nutrient limitation of cultured algae.

Physical factors driving intertidal macroalgae distribution: physiological stress of a dominant fucoid at its southern limit

Climate change is driving species range shifts worldwide. However, physiological responses related to distributional changes are not fully understood. Oceanographers have reported an increase in ocean temperature in the northwest Iberian Peninsula that is potentially related to the decline in some cold-temperate intertidal macroalgae in the Cantabrian Sea, namely Fucus serratus. Low tide stress could also play a role in this decline. We performed one mensurative (in situ) and two manipulative (in culture) experiments designed to evaluate the interactive effects of some physical factors. The first experiment analysed field response to low tide stress in marginal (mid-Cantabrian Sea and northern Portugal) versus central (Galicia) populations of F. serratus. Then a second experiment was performed that utilized either harsh or mild summer conditions of atmospheric temperature, irradiance, humidity, and wind velocity to compare the responses of individuals from one marginal and one central population to low tide stress. Finally, the combined effect of sea temperature and the other factors was evaluated to detect interactive effects. Changes in frond growth, maximal photosynthetic quantum yield (Fv/Fm), temperature, and desiccation were found. Three additive factors (solar irradiation, ocean and air temperatures) were found to drive F. serratus distribution, except under mildly humid conditions that ameliorated atmospheric thermal stress (two additive factors). Mid-Cantabrian Sea temperatures have recently increased, reaching the inhibitory levels suggested in this study of F. serratus. We also expect an additive secondary contribution of low tide stress to this species decline. On the northern Portugal coast, ocean warming plus low tide stress has not reached this species’ inhibition threshold. No significant differential responses attributed to the population of origin were found. Mechanistic approaches that are designed to analyse the interactive effects of physical stressors may improve the levels of confidence in predicted range shifts of species.

Carotenoid composition in Rhodophyta: insights into xanthophyll regulation in Corallina elongata

Macroalgae possess several photoprotection mechanisms, including xanthophyll cycles. Among these cycles, the VAZ (violaxanthin–anteraxanthin–zeaxanthin) cycle controls the interconversion of epoxidated xanthophyll (V) to de-epoxidated forms anteraxanthin and zeaxanthin. This conversion modulates the amount of excess energy that it is dissipated as heat. Presence of functional xanthophyll cycles in Rhodophyta is controversial. In this work we investigated the presence of xanthophyll cycles in red algae. Carotenoid composition of 13 red macroalgae collected in northern Spain was studied by HPLC. This screening showed that carotenoid composition is a conservative trade in rhodophytes, with very few exceptions to the general pattern formed by β-carotene and one to three xanthophylls: anteraxanthin, zeaxanthin or lutein. Corallina elongata and Jania rubens were the only algae containing anteraxanthin as the main xanthophyll. The first species was selected to study whether any truncated xanthophyll cycle between anteraxanthin and zeaxanthin could operate under photoinhibitory conditions. Upon illumination, xanthophyll composition remained stable, without any short-term light-induced de-epoxidation of anteraxanthin to zeaxanthin. However, an inverse relation between both xanthophylls was found. In addition, a seasonal trend of changes in anteraxanthin and zeaxanthin was observed when pigment composition was studied in field samples over the course of a year. We concluded that these variations were more likely due to differential rates of synthesis and degradation of xanthophylls than to the operation of a xanthophyll cycle. The great amount of anteraxanthin in C. elongata may play an structural stability role on light-harvesting complexes.

INORGANIC NITROGEN AND PHOSPHORUS UPTAKE KINETICS IN PALMARIA PALMATA (RHODOPHYTA) 1

The N and P uptake responses were studied in a northern Spanish population of the edible red seaweed Palmaria palmata (Linnaeus) Kuntze. The fronds were incubated at different concentrations, and the nutrient depletion in the medium was measured at successive times to calculate uptake rates. Palmaria palmata uptake response was biphasic and nonsaturable for inorganic P. This would allow the species to exploit transient pulses of high P concentration in natural and fertilized conditions. Such a response is a common feature of algae avoiding nutrient deficiency. At average concentrations measured in the ocean, the response was nonsaturable for inorganic N sources, except for ammonium in autumn and winter when it is not the major N source. In contrast to the general rule of ammonium being taken at a higher rate than nitrate, we found similar affinity for both nutrients corresponding to the minor role of ammonium as N source for field populations over the year.

Physiological response of fucoid algae to environmental stress: comparing range centre and southern populations

Climate change has led to alterations in assemblage composition. Species of temperate macroalgae at their southern limits in the Iberian Peninsula have shown shifts in geographical range and a decline in abundance ultimately related to climate, but with the proximate factors largely unknown. We performed manipulative experiments to compare physiological responses of Fucus vesiculosus and Fucus spiralis from Portugal and Wales (UK), representing, respectively, southern and central areas of their distribution, to different intensities of solar radiation and different air temperatures. Following exposure to stressful emerged conditions, Portuguese and Welsh individuals of both fucoid species showed increased frond temperature, high desiccation levels and reduced photophysiological performance that was evident even after a 16 h recovery period, with light and temperature acting in an additive, not an interactive, manner. The level of physiological decline was influenced by geographical origin of populations and species identity, with algae from the south and those living higher on the shore coping better with stressful conditions. The negative effect of summer conditions on photophysiology may contribute to changes in fucoid abundance and distribution in southern Europe. Our results emphasise how physiological performance of geographically distinct populations can differ, which is particularly relevant when predicting responses to climate change.

CHANGES IN NUTRIENT CONTENT OF PALMARIA PALMATA IN RESPONSE TO VARIABLE LIGHT AND UPWELLING IN NORTHERN SPAIN1

Light has been identified as one of the main factors affecting seaweed ecophysiology. We investigated the dependence of nutrient metabolism on sun and shade light conditions and whether episodes of upwelling of nutrient‐rich subsuperficial water could reduce the summer nutrient limitation driving physiological changes in Palmaria palmata (L.) Kuntze. We measured the major nutrient pools, photosynthetic pigments, and light curves, under sun and shade conditions during a summer period when one upwelling was recorded. The redundancy analysis (RDA) produced two clear groups: sun‐ and shade‐acclimated algae. Light was the major predictive factor. Sun‐acclimated algae exhibited higher carbon (C) and lower nitrogen (N) and phosphorus (P) content in association with the storage of floridoside (main C reserve) to benefit from higher irradiance (under nutrient limitation). Among N pools, N reserves (phycoerythrin, nitrate) were a lower proportion of the total N in sun‐acclimated algae, suggesting their degradation to fulfill the N demands of the cell. The orthophosphate content was also lower in sun‐acclimated algae, indicating its utilization as a nutrient reserve. In contrast, N within cell walls and membranes and chl a contributed to a similar proportion of the total N in sun‐ and shade‐acclimated algae, suggesting a response to sustain cell integrity. Transient high nutrient concentration due to the upwelling was unrelated to the nutrient content of the thallus. The storage of C as floridoside from high light exposure was shown to be the driving force for the metabolic adjustment of P. palmata at the end of summer before the onset of dormancy.

The ‘golden kelp’ Laminaria ochroleuca under global change: integrating multiple eco-physiological responses with species distribution models

1.The loss of marine foundation species, in particular kelps at temperate latitudes, has been linked to climatic drivers and co-occurring human perturbations. Ocean temperature and nutrients typically co-vary over local and regional scales and play a crucial role on kelp dynamics. Examining their independent and interactive effects on kelp physiological performance is essential to understand and predict patterns of kelp distribution, particularly under scenarios of global change. 2.Crossed combinations of ocean temperatures and availability of nutrients were experimentally tested on juveniles of the ‘golden kelp’, Laminaria ochroleuca, from the northwestern Iberian Peninsula. Eco-physiological responses included: survival, growth and total N content. Results were embedded into a Species Distribution Model (SDM), which relates presence records and climatic and non-climatic data to forecast distribution patterns of L. ochroleuca under different climate change scenarios. 3.Temperatures above 24.6 °C were lethal irrespective of nutrients. Optimal growth of juvenile sporophytes occurred between 12 °C and 18 °C and no nutrient limitation. The SDM, where ocean temperature was the main predictor of kelp distribution in line with temperature thresholds given by eco-physiological responses, suggests a future expansion towards northern latitudes and a retreat from the southern limit/boundary of the current distribution. 4.Synthesis. Range-shifting of the golden kelp can have severe ecological impacts at regional and local scales. The expansion or retraction of the species along the European coast seems to be modulated mainly by temperature, but nutrient availability would be key to maintain optimal physiological performance. Our work highlights that the combination of empirical and modelling approaches is accessible to researchers and crucial to building more robust predictions of ecological and biogeographic responses of habitat-forming species to forecasted environmental change. This article is protected by copyright. All rights reserved.

Physiological responses to variations in grazing and light conditions in native and invasive fucoids

Poor physiological acclimatization to climate change has led to shifts in the distributional ranges of various species and to biodiversity loss. However, evidence also suggests the relevance of non-climatic physical factors, such as light, and biotic factors, which may act in interactive or additive way. We used a mechanistic approach to evaluate the ecophysiological responses of four seaweed species (three dominant intertidal fucoids, Fucus serratus, Ascophyllum nodosum, Bifurcaria bifurcata, and the invasive Sargassum muticum) to different conditions of grazing, light irradiance and ultraviolet (UV) radiation. We performed a large-scale mesocosm experiment with a total of 800 individual thalli of macroalgae. The factorial experimental design included major algal traits, photoacclimation, nutrient stoichiometry and chemical defence as response variables. Few significant effects of the factors acting alone or in combination were observed, suggesting a good capacity for acclimatization in all four species. The significant effects were generally additive and there were no potentially deleterious synergistic effects between factors. Fucus serratus, a species currently undergoing a drastic contraction of its southern distribution limit in Europe, was the most strongly affected species, showing overall lower photosynthetic efficiency than the other species. The growth rate of F. serratus decreased when UV radiation was filtered out, but only in the presence of grazers. Moreover, more individuals of this species tended to reach maturity in the absence of grazers, and the nitrogen content of tissues decreased under full-spectrum light. Only the phlorotannin content of tissues of B. bifurcata and of exudates of A. nodosum, both slow-growing species, were positively affected by respectively removal of UVB radiation and the presence of grazers. The findings for S. muticum, a well-established invasive seaweed across European coasts, suggested similar physiological response of this fast-growing species to different levels of grazing activity and light quality/intensity. As expected, this species grew faster than the other species. Bifurcaria bifurcata and A. nodosum only showed minor effects of light quality and grazing on phlorotannins content, which suggests good resistance of these two long-lived species to the experimental conditions. Mechanistic approaches that are designed to analyse interactive effects of physical and biotic factors provide an understanding of physiological responses of species and help to improve the confidence of predictive distribution models.