Seaweed micropropagation techniques and their potentials: an overview

The seaweed industry worldwide uses 7.5–8.0 million tonnes of wet seaweeds annually with a majority of it derived from cultivated farms, as the demand for seaweed based-products exceeds the supply of seaweed raw material from natural stocks. The main advantage of cultivation is that it not only obviates overexploitation of natural populations but also facilitates the selection of germplasm with desired traits. To enhance the economic prospects of seaweed cultivation, varied practices, such as simple and cost effective cultivation methods, use of select germplasm as seed stock coupled with good farm management practices, etc., are adopted. Nevertheless, in vitro cell culture techniques have also been employed as they facilitate development and propagation of genotypes of commercial importance. There are more than 85 species of seaweeds for which tissue culture aspects have been reported. Although the initial aim of these techniques focuses mostly on genetic improvement and clonal propagation of seaweeds for mariculture, recently the scope of these techniques has been extended for use in bioprocess technology for production of high value chemicals of immense importance in the pharmaceutical and nutraceutical sectors. Recently, there has been a phenomenal interest in intensifying seaweed tissue and cell culture research to maximize the add-on value of seaweed resources. This paper deals with the status of seaweed micropropagation techniques and their applications in the context of the marine biotech industry. Further, it also provides an analysis of the problems to be resolved for removing the barriers that are impeding the true realization of potentials offered by these techniques for sustainable development and utilization of seaweed resources.     

Drag reduction in wave-swept macroalgae: Alternative strategies and new predictions

? Premise of the study: Intertidal macroalgae must resist extreme hydrodynamic forces imposed by crashing waves. How does frond flexibility mitigate drag, and how does flexibility affect predictions of drag and dislodgement in the field? ? Methods: We characterized flexible reconfiguration of six seaweed species in a recirculating water flume, documenting both shape change and area reduction as fronds reorient. We then used a high-speed gravity-accelerated water flume to test our ability to predict drag under waves based on extrapolations of drag recorded at slower speeds. We compared dislodgement forces to drag forces predicted from slow- and high-speed data to generate new predictions of survivorship and maximum sustainable frond size along wave-swept shores. ? Key results: Bladed algae were generally "shape changers", limiting drag by reducing drag coefficients, whereas the branched alga Calliarthron was an "area reducer", limiting drag by reducing projected area in flow. Drag predictions often underestimated actual drag measurements at high speeds, suggesting that slow-speed data may not reflect the performance of flexible seaweeds under breaking waves. Several seaweeds were predicted to dislodge at similar combinations of velocity and frond size, suggesting common scaling factors of dislodgement strength and drag. ? Conclusions: Changing shape and reducing projected area in flow are two distinct strategies employed by flexible seaweeds to resist drag. Flexible reconfiguration contributes to the uncertainty of drag extrapolation, and researchers should use caution when predicting drag and dislodgement of seaweeds in the field.     

Ultraviolet radiation shapes seaweed communities

Stratospheric ozone depletion and the concomitant increase in irradiance of ultraviolet-B radiation (UVB) at the earth’s surface represent major threats to terrestrial and aquatic ecosystems. In costal rocky shore environments, seaweeds constitute a group of organisms of particular significance to ecosystem function. Thus, impairment of seaweed performance by UVB-exposure may result in severe changes in the functioning of coastal ecosystems. Here we present our view on how UVB radiation affects seaweed physiology and ecology and, thus, shapes the coastal environment by affecting the spatial, species and functional structure of seaweed communities.     

Finite element modeling of shell shape in the freshwater turtle Pseudemys concinna reveals a trade‐off between mechanical strength and hydrodynamic efficiency

Aquatic species can experience different selective pressures on morphology in different flow regimes. Species inhabiting lotic regimes often adapt to these conditions by evolving low‐drag (i.e., streamlined) morphologies that reduce the likelihood of dislodgment or displacement. However, hydrodynamic factors are not the only selective pressures influencing organismal morphology and shapes well suited to flow conditions may compromise performance in other roles. We investigated the possibility of morphological trade‐offs in the turtle Pseudemys concinna. Individuals living in lotic environments have flatter, more streamlined shells than those living in lentic environments; however, this flatter shape may also make the shells less capable of resisting predator‐induced loads. We tested the idea that “lotic” shell shapes are weaker than “lentic” shell shapes, concomitantly examining effects of sex. Geometric morphometric data were used to transform an existing finite element shell model into a series of models corresponding to the shapes of individual turtles. Models were assigned identical material properties and loaded under identical conditions, and the stresses produced by a series of eight loads were extracted to describe the strength of the shells. “Lotic” shell shapes produced significantly higher stresses than “lentic” shell shapes, indicating that the former is weaker than the latter. Females had significantly stronger shell shapes than males, although these differences were less consistent than differences between flow regimes. We conclude that, despite the potential for many‐to‐one mapping of shell shape onto strength, P. concinna experiences a trade‐off in shell shape between hydrodynamic and mechanical performance. This trade‐off may be evident in many other turtle species or any other aquatic species that also depend on a shell for defense. However, evolution of body size may provide an avenue of escape from this trade‐off in some cases, as changes in size can drastically affect mechanical performance while having little effect on hydrodynamic performance. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Acclimation of bloom‐forming and perennial seaweeds to elevated pCO2 conserved across levels of environmental complexity

Macroalgae contribute approximately 15% of the primary productivity in coastal marine ecosystems, fix up to 27.4 Tg of carbon per year, and provide important structural components for life in coastal waters. Despite this ecological and commercial importance, direct measurements and comparisons of the short‐term responses to elevated pCO₂ in seaweeds with different life‐history strategies are scarce. Here, we cultured several seaweed species (bloom forming/nonbloom forming/perennial/annual) in the laboratory, in tanks in an indoor mesocosm facility, and in coastal mesocosms under pCO₂ levels ranging from 400 to 2,000 μatm. We find that, across all scales of the experimental setup, ephemeral species of the genus Ulva increase their photosynthesis and growth rates in response to elevated pCO₂ the most, whereas longer‐lived perennial species show a smaller increase or a decrease. These differences in short‐term growth and photosynthesis rates are likely to give bloom‐forming green seaweeds a competitive advantage in mixed communities, and our results thus suggest that coastal seaweed assemblages in eutrophic waters may undergo an initial shift toward communities dominated by bloom‐forming, short‐lived seaweeds.     

Review: Seaweed tissue culture as applied to biotechnology: Problems,achievements and prospects

Advances have been made in cell and tissue culture of seaweeds to define a unique branch of in vitro techniques; however, they are lagging far behind those of land plants and have limited applications. Explants can be cultivated axenically in enriched or artificial seawater culture media, and regeneration and even callus formation are achieved. In this state of the art technique, seaweed tissue culture may be already useful for certain biotechnological applications, such as clonal propagation of seed material for mariculture. Nevertheless, the absolute control of growth and development as it is exerted in higher plant tissue culture is lacking, and it is required for more complex biotechnological applications in seaweeds. Definitively, we need appropriate cells (competent cells) to induce growth with the most effective chemical regulators in culture medium adjusted towards the addition of carbon sources. Still, free cells and protoplast isolation and regeneration in marine seaweeds constitute the most developed topic in seaweed tissue culture. The regulation of growth and development of seaweed free cell and protoplast cultures may sustain a purposeful use of techniques in the era of genomic applications.     

Prospects and challenges for industrial production of seaweed bioactives

Large‐scale seaweed cultivation has been instrumental in globalizing the seaweed industry since the 1950s. The domestication of seaweed cultivars (begun in the 1940s) ended the reliance on natural cycles of raw material availability for some species, with efforts driven by consumer demands that far exceeded the available supplies. Currently, seaweed cultivation is unrivaled in mariculture with 94% of annual seaweed biomass utilized globally being derived from cultivated sources. In the last decade, research has confirmed seaweeds as rich sources of potentially valuable, health‐promoting compounds. Most existing seaweed cultivars and current cultivation techniques have been developed for producing commoditized biomass, and may not necessarily be optimized for the production of valuable bioactive compounds. The future of the seaweed industry will include the development of high value markets for functional foods, cosmeceuticals, nutraceuticals, and pharmaceuticals. Entry into these markets will require a level of standardization, efficacy, and traceability that has not previously been demanded of seaweed products. Both internal concentrations and composition of bioactive compounds can fluctuate seasonally, geographically, bathymetrically, and according to genetic variability even within individual species, especially where life history stages can be important. History shows that successful expansion of seaweed products into new markets requires the cultivation of domesticated seaweed cultivars. Demands of an evolving new industry based upon efficacy and standardization will require the selection of improved cultivars, the domestication of new species, and a refinement of existing cultivation techniques to improve quality control and traceability of products.     

183 The Environmental Impacts of Harvesting Beach‐Cast Seaweeds

Until recently the commercial collection of beach‐cast seaweeds in New Zealand was prohibited but the legislation has recently been amended to allow permitting of this activity. This review collates existing information on the role of beach‐cast seaweed in coastal ecosystems to describe the nature and extent of the effects that commercial removals of beach cast seaweed may have on the marine environment. It outlines the amount of beach‐cast seaweed available for harvest in New Zealand and the fate of seaweed when not collected; reviewing current information on the importance of beach‐cast seaweeds and its inhabitants on: feeding and nesting shorebirds, and nutritional contribution of seaweed inhabitants to nearshore coastal ecosystems when seaweed is washed back into the sea It also identifies key research gaps related to any environmental impacts that the removal of beach cast seaweed might have.     

Genetic improvement of macroalgae: status to date and needs for the future

Marine macroalgae (seaweeds) consist of a diverse range of species with diverse morphology and bioactive properties. The potential for genetic improvement to increase the value of the production of such molecules and other economically important traits is likely to be high as procedures for cultivation of many species are well established, genetic diversity is often high, sexual propagation is often possible and seaweeds generally have a short generation time that allows rapid transition from one selected generation to the next. The need for genetic improvement has been voiced by industry for many years. Despite this, there is little published research describing seaweed genetic improvement methodology, results or impact. This review draws on knowledge from other groups of species and describes options, benefits and knowledge needed for the genetic improvement of macroalgae. In conclusion, the review highlights how a well-designed genetic improvement program, which targets one or a few select economically valuable traits whilst limiting inbreeding, could provide potentially large benefits for industries reliant on the culture of seaweeds. Progress will, to a large extent, rely on the magnitude of genetic variation for the trait in question.     

THE EVOLVABILITY OF GROWTH FORM IN A CLONAL SEAWEED

Although modular construction is considered the key to adaptive growth or growth‐form plasticity in sessile taxa (e.g., plants, seaweeds and colonial invertebrates), the serial expression of genes in morphogenesis may compromise its evolutionary potential if growth forms emerge as integrated wholes from module iteration. To explore the evolvability of growth form in the red seaweed, Asparagopsis armata, we estimated genetic variances, covariances, and cross‐environment correlations for principal components of growth‐form variation in contrasting light environments. We compared variance–covariance matrices across environments to test environmental effects on heritable variation and examined the potential for evolutionary change in the direction of plastic responses to light. Our results suggest that growth form in Asparagopsis may constitute only a single genetic entity whose plasticity affords only limited evolutionary potential. We argue that morphological integration arising from modular construction may constrain the evolvability of growth form in Asparagopsis, emphasizing the critical distinction between genetic and morphological modularity in this and other modular taxa.     

SELF‐THINNING AND SIZE INEQUALITY DYNAMICS IN A CLONAL SEAWEED (SARGASSUM LAPAZEANUM,PHAEOPHYCEAE)1

Fronds of clonal seaweeds with extensive holdfasts relative to frond size are known not to self‐thin during growth, even in crowded stands. We tested whether frond self‐thinning would occur for such a seaweed since these traits are more similar to those of unitary seaweeds, which do self‐thin in crowded conditions. We used Sargassum lapazeanum Setch. et N. L. Gardner (Fucales, Phaeophyceae) from the Pacific coast of Mexico, for which we first confirmed its clonal nature by performing a regeneration experiment in culture tanks. During the growth season (winter to late spring), S. lapazeanum stand biomass increased, while frond density and size inequality (Gini coefficient for frond biomass) decreased. These results indicate that self‐thinning occurred at the frond level. We propose a conceptual model for frond dynamics for clonal seaweeds in general. In stands of clonal species with small fronds and relatively extensive holdfasts (particularly when holdfasts are perennial), frond dynamics would be determined mostly by intraclonal regulation, which seems to prevent excessive crowding from occurring. Such species display a positive biomass–density relationship during the growth season. On the contrary, in stands of clonal species with large fronds relative to holdfast size, frond dynamics would be determined mostly by interactions among genets. For such species, self‐thinning may be detected at the frond level in crowded stands, resulting in a negative biomass–density relationship during growth.     

Comment on ‘Island biogeography: patterns of marine shallow‐water organisms’ by Hachich et al., Journal of Biogeography (2015)

In a recent article, Hachich et al. (2015, Journal of Biogeography, 42 , 1871–1882) studied the large‐scale biogeographical patterns of the species–area, species–island age and species–isolation relationships associated with marine shallow‐water groups (reef fish, gastropods and seaweeds) from 11 Atlantic archipelagos. We here express our concerns regarding the data accuracy used to compute the different models that tested the null hypothesis of species richness being independent of the selected variables. In our commentary, we focus mainly on the use of out‐of‐date checklists of gastropod and seaweed species from different archipelagos, but we also point out inaccuracies in some island age estimates and explain our disagreement with the use of the 200 m depth limit for the shallow‐water gastropods and seaweeds.     

The use of Gracilaria tikvahiae (Gracilariales,Rhodophyta) as a model system to understand the nitrogen nutrition of cultured seaweeds

Seaweeds have physiological mechanisms to acquire, utilize, and store various forms of nitrogen in environments where nitrogen levels vary tremendously in space and time. Knowledge of the nitrogen relationships of seaweeds is required for the development of successful seaweed mariculture. For example, it would seem at first that continuous nitrogen enrichment would be desirable in such systems because maximal seaweed yields are possible only when growth is not nitrogen-limited. Yet such fertilization is wasteful and can result in yield reductions due to the enhancement of epiphyte growth. Because most seaweeds can rapidly taken up high concentrations of nitrogen, far in excess of what is required for current growth demands, enrichments are needed only when internal nitrogen concentrations fall to near the critical level (i.e., the minimal tissue concentration of nitrogen required for maximal growth). Nutrients are best applied at brief pulses of high nitrogen concentrations.Dedicated to the memory of Bud Brinkhuis, friend and colleague     

Variation in the impact of non‐native seaweeds along gradients of habitat degradation: a meta‐analysis and an experimental test

Biological invasions are acknowledged among the main drivers of global changes in biodiversity. Despite compelling evidence of species interactions being strongly regulated by environmental conditions, there is a dearth of studies investigating how the effects of non‐native species vary among areas exposed to different anthropogenic pressures. Focusing on marine macroalgae, we performed a meta‐analysis to test whether and how the direction and magnitude of their effects on resident communities and species varies in relation to cumulative anthropogenic impact levels. The relationship between human impact levels and non‐native species impact intensity emerged only for a reduced subset of the response variables examined. Yet, there was a trend for the effects of non‐native species on community biomass and abundance and on species abundance to become less negative at heavily impacted sites. By contrast, the magnitude of negative effects of seaweed on community evenness tended to increase with human impact levels. The hypothesis of decreasing severity of invader’ impacts along a gradient of habitat degradation was also tested experimentally at a regional scale by comparing the effects of the removal of non‐native alga, Caulerpa cylindracea, on resident assemblages among rocky reefs exposed to different anthropogenic pressures. Assemblages at urban and pristine site did not differ when invaded, but did so when C. cylindracea was removed. Our results suggest that, despite the generally weak relationship between human impacts levels and non‐native species impacts, more negative impacts can be expected in less stressful environments (i.e. less degraded or pristine sites), where competitive interactions are presumably the driving force structuring resident communities. Implementing strategies for controlling the establishment of non‐native seaweeds should be, thus, considered a priority for preserving biodiversity in relatively pristine areas. On the other hand, control of invaders at degraded sites could be warranted to lessen their role as propagule sources. Synthesis Local anthropogenic stressors that severely alter biotic and abiotic conditions may underpin context‐dependency in the impacts of biological invasions. We used a meta‐analysis and an experimental test to examine the relationship between cumulative human impacts and ecological impact of non‐native seaweeds on resident assemblages. Our results suggest that more negative impacts of non‐native seaweeds on the abundance and biomass of resident assemblages can be expected in less degraded or pristine sites. Possibly, stronger impacts prevail at pristine sites, where assemblages are mainly structured by biotic interactions. Hence, management efforts should be mostly directed to prevent the establishment and spread of non‐native seaweeds in pristine areas. On the other hand, weak, but positive effects of seaweeds at the most degraded sites add to the ongoing debate on the role of non‐native species in rehabilitation plans.     

Ontogenetic modulation of branch size,shape, and biomechanics produces diversity across habitats in the Bursera simaruba clade of tropical trees

Organismal size and shape inseparably interact with tissue biomechanical properties. It is therefore essential to understand how size, shape, and biomechanics interact in ontogeny to produce morphological diversity. We estimated within species branch length‐diameter allometries and reconstructed the rates of ontogenetic change along the stem in mechanical properties across the simaruba clade in the tropical tree genus Bursera, measuring 376 segments from 97 branches in nine species in neotropical dry to rain forest. In general, species with stiffer materials had longer, thinner branches, which became stiffer more quickly in ontogeny than their counterparts with more flexible materials. We found a trend from short stature and flexible tissues to tall statures and stiff tissues across an environmental gradient of increasing water availability, likely reflecting a water storage–mechanical support tradeoff. Ontogenetic variation in size, shape, and mechanics results in diversity of habits, for example, rapid length extension, sluggish diameter expansion, and flexible tissues results in a liana, as in Bursera instabilis. Even species of similar habit exhibited notable changes in tissue mechanical properties with increasing size, illustrating the inseparable relationship between organismal proportions and their tissue mechanics in the ontogeny and evolution of morphological diversity.     

A sixth‐level habitat cascade increases biodiversity in an intertidal estuary

Many studies have documented habitat cascades where two co‐occurring habitat‐forming species control biodiversity. However, more than two habitat‐formers could theoretically co‐occur. We here documented a sixth‐level habitat cascade from the Avon‐Heathcote Estuary, New Zealand, by correlating counts of attached inhabitants to the size and accumulated biomass of their biogenic hosts. These data revealed predictable sequences of habitat‐formation (=attachment space). First, the bivalve Austrovenus provided habitat for green seaweeds (Ulva) that provided habitat for trochid snails in a typical estuarine habitat cascade. However, the trochids also provided habitat for the nonnative bryozoan Conopeum that provided habitat for the red seaweed Gigartina that provided habitat for more trochids, thereby resetting the sequence of the habitat cascade, theoretically in perpetuity. Austrovenus is here the basal habitat‐former that controls this “long” cascade. The strength of facilitation increased with seaweed frond size, accumulated seaweed biomass, accumulated shell biomass but less with shell size. We also found that Ulva attached to all habitat‐formers, trochids attached to Ulva and Gigartina, and Conopeum and Gigartina predominately attached to trochids. These “affinities” for different habitat‐forming species probably reflect species‐specific traits of juveniles and adults. Finally, manipulative experiments confirmed that the amount of seaweed and trochids was important and consistent regulators of the habitat cascade in different estuarine environments. We also interpreted this cascade as a habitat‐formation network that describes the likelihood of an inhabitant being found attached to a specific habitat‐former. We conclude that the strength of the cascade increased with the amount of higher‐order habitat‐formers, with differences in form and function between higher and lower‐order habitat‐formers, and with the affinity of inhabitants for higher‐order habitat‐formers. We suggest that long habitat cascades are common where species traits allow for physical attachment to other species, such as in marine benthic systems and old forest.     

Effects of invasive seaweeds on feeding preference and performance of a keystone Mediterranean herbivore

The consequences of invasive species on ecosystem processes and ecological interactions remain poorly understood. Predator–prey interactions are fundamental in shaping species evolution and community structure and can be strongly modified by species introductions. To fully understand the ecological effects of invasive species on trophic linkages it is important to characterize novel interactions between native predators and exotic prey and to identify the impacts of invasive species on the performance of native predators. Although seaweed invasions are a growing global concern, our understanding of invasive algae—herbivore interactions is still very limited. We used a series of feeding experiments between a native herbivore and four invasive algae in the Mediterranean Sea to examine the potential of native sea urchins to consume invasive seaweeds and the impacts of invasive seaweed on herbivore performance. We found that three of the four invasive species examined are avoided by native herbivores, and that feeding behaviour in sea urchins is not driven by plant nutritional quality. On the other hand, Caulerpa racemosa is readily consumed by sea urchins, but may escape enemy control by reducing their performance. Recognizing the negative impacts of C. racemosa on herbivore performance has highlighted an enemy escape mechanism that contributes to explaining how this widespread invasive alga, which is preferred and consumed by herbivores, is not eradicated by grazing in the field. Furthermore, given the ecological and economic importance of sea urchins, negative impacts of invasive seaweeds on their performance could have dramatic effects on ecosystem function and services, and should be accounted for in sea urchin population management strategies.     

IMPORTANCE OF MACRO‐ VERSUS MICROSTRUCTURE IN MODULATING LIGHT LEVELS INSIDE CORAL COLONIES1

Adjusting the light exposure and capture of their symbiotic photosynthetic dinoflagellates (genus Symbiodinium Freud.) is central to the success of reef‐building corals (order Scleractinia) across high spatio‐temporal variation in the light environment of coral reefs. We tested the hypothesis that optical properties of tissues in some coral species can provide light management at the tissue scale comparable to light modulation by colony architecture in other species. We compared within‐tissue scalar irradiance in two coral species from the same light habitat but with contrasting colony growth forms: branching Stylophora pistillata and massive Lobophyllia corymbosa. Scalar irradiance at the level of the symbionts (2 mm into the coral tissues) were <10% of ambient irradiance and nearly identical for the two species, despite substantially different light environments at the tissue surface. In S. pistillata, light attenuation (90% relative to ambient) was observed predominantly at the colony level as a result of branch‐to‐branch self‐shading, while in L. corymbosa, near‐complete light attenuation (97% relative to ambient) was occurring due to tissue optical properties. The latter could be explained partly by differences in photosynthetic pigment content in the symbiont cells and pigmentation in the coral host tissue. Our results demonstrate that different strategies of light modulation at colony, polyp, and cellular levels by contrasting morphologies are equally effective in achieving favorable irradiances at the level of coral photosymbionts.     

Seaweed production: overview of the global state of exploitation,farming and emerging research activity

ABSTRACT

The use of seaweeds has a long history, as does the cultivation of a select and relatively small group of species. This review presents several aspects of seaweed production, such as an update on the volumes of seaweeds produced globally by both extraction from natural beds and cultivation. We discuss uses, production trends and economic analysis. We also focus on what is viewed as the huge potential for growing industrial-scale volumes of seaweeds to provide sufficient, sustainable biomass to be processed into a multitude of products to benefit humankind. The biorefinery approach is proposed as a sustainable strategy to achieve this goal. There are many different technologies available to produce seaweed, but optimization and more efficient developments are still required. We conclude that there are some fundamental and very significant hurdles yet to overcome in order to achieve the potential contributions that seaweed cultivation may provide the world. There are critical aspects, such as improving the value of seaweed biomass, along with a proper consideration of the ecosystem services that seaweed farming can provide, e.g. a reduction in coastal nutrient loads. Additional considerations are environmental risks associated with climate change, pathogens, epibionts and grazers, as well as the preservation of the genetic diversity of cultivated seaweeds. Importantly, we provide an outline for future needs in the anticipation that phycologists around the world will rise to the challenge, such that the potential to be derived from seaweed biomass becomes a reality.     

High CO2 enhances the competitive strength of seaweeds over corals

Space competition between corals and seaweeds is an important ecological process underlying coral-reef dynamics. Processes promoting seaweed growth and survival, such as herbivore overfishing and eutrophication, can lead to local reef degradation. Here, we present the case that increasing concentrations of atmospheric CO(2) may be an additional process driving a shift from corals to seaweeds on reefs. Coral (Acropora intermedia) mortality in contact with a common coral-reef seaweed (Lobophora papenfussii) increased two- to threefold between background CO(2) (400 ppm) and highest level projected for late 21st century (1140 ppm). The strong interaction between CO(2) and seaweeds on coral mortality was most likely attributable to a chemical competitive mechanism, as control corals with algal mimics showed no mortality. Our results suggest that coral (Acropora) reefs may become increasingly susceptible to seaweed proliferation under ocean acidification, and processes regulating algal abundance (e.g. herbivory) will play an increasingly important role in maintaining coral abundance.