Macrocystis Mariculture in Chile : Growth Performance of Heterosis
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Erschienen in: Journal of Applied Phycology ; 23 (2011), 5. - S. 819-825 https://dx.doi.org/10.1007/s10811-010-9581-z Macrocystis mariculture in Chile: growth performance of heterosis genotype constructs under field conditions Renato Westermeier & David J. Patiño & Pedro Murúa & Dieter G. Müller Abstract Recent progress in Macrocystis mariculture is Introduction based on clonal stock cultures of gametophyte parents. Batches of up to 105 genetically identical sporophyte Macrocystis (giant kelp) is an important natural resource in seedlings can be produced at any time in the laboratory the coastal waters of Chile. Dried material is used for and explanted in the field for production of biomass. Sexual alginate production, while fresh fronds are harvested in crosses of selected Macrocystis pyrifera gametophyte large quantities as food for mariculture of herbivorous high- parents of different geographic origin along the coast of value mollusks (abalone, Haliotis rufescens and H. discus Chile showed heterosis and produced sporophyte batches hannai). Intense harvesting pressure on natural kelp beds with superior growth performance. Starting from zygotes, has led to over-exploitation damage (Vásquez 2008). This after 10 weeks in the laboratory and 5 months in the sea, situation calls for efforts to supplement the available kelp our best hybrid genotypes grew up to 11 kg fresh weight biomass by laboratory-based culture methods. per frond, which corresponds to 66 kg m 1 of line in a Traditional mariculture techniques make use of meio- commercial mariculture installation. In contrast, average spores from natural populations, which are collected and yields of 14.4 and 22 kg m 1 are reported in the literature manipulated to settle on ropes in laboratory tanks. The for traditional methods. Additional experiments, including resulting artificial substrates with juvenile sporophytes are inter-specific crosses M. pyrifera × M. integrifolia and their explanted to mariculture installations in the sea (Gutierrez performance in different climate zones of Chile, confirm et al. 2006; Macchiavello et al. 2010). This method has that heterosis is a powerful tool for crop improvement in significant disadvantages, mainly the seasonal restriction Macrocystis. It opens the possibility to construct tailor-made and quality of natural spore supply and problems to obtain heterosis genotypes with maximum productivity and/or other proper inoculation densities. Furthermore, propagules of desired properties for any given locality. epiflora and epifauna introduced with natural spores cause serious fouling problems. Growth rates and harvest biomass Keywords Biomass . Chile . Cross-breeding . Heterosis . per area in such plantations are not satisfactory and subject Macrocystis . Mariculture to erratic variations. These inherent deficits of traditional Macrocystis farming call for fundamentally different novel approaches in Macrocystis mariculture management. Inno- vations like buildup of permanent genetic stock, standard- * : : R. Westermeier ( ) D. J. Patiño P. Murúa ized seedling production and selection of favourable Instituto de Acuicultura, Universidad Austral de Chile, Campus Puerto Montt, Casilla 1327, genotypes are now well under way (Westermeier et al. Puerto Montt, Chile 2010). These features fulfil the status of “domestication” e mail: [email protected] for the kelp Macrocystis, which can be defined as “the process of hereditary reorganization of wild animals and D. G. Müller Fachbereich Biologie der Universität Konstanz, plants into forms more accommodating to the interests of 78457 Constance, Germany people” (Anonymous 2010). Konstanzer Online-Publikations-System (KOPS) URL: http://nbn-resolving.de/urn:nbn:de:bsz:352-0-306974 820 The life history of Macrocystis and related kelps is well population senescence may be responsible for spontaneous studied (Graham et al. 2007). As members of the order population oscillations in Californian Macrocystis beds. Laminariales, they exhibit a biphasic cycle, which alternates From these considerations, it is evident that the Macro between a macroscopic sporophyte and a microscopic cystis inoculants hitherto used for mariculture are likely to gametophyte. Sporophytes are diploid and fixed to the be handicapped by inbreeding depression. Both the tradi- substrate by a multicellular holdfast. Upright growth of fronds tional rope-seeding method as well as the use of one pair of is accomplished by apical meristems which differentiate into genetically related gametophyte clones for laboratory metre-sized stipes and lateral blades for photosynthesis. seedling production are subject to the burden of inbreeding Sporophytes enter reproduction by forming sori on fertile effects. In consequence, outcrossing must be expected to be blades, where numerous superficial thallus cells undergo a promising way to improve the growth characteristics of meiosis. Huge numbers of sporangia appear, each releasing 32 laboratory-produced Macrocystis seedlings. motile meiospores. Upon fixation to the substrate, spores In order to evaluate this possibility, Westermeier et al. develop into haploid, few-celled uni-seriate gametophytes. (2010) carried out a systematic cross-breeding programme Due to genotypic sex determination, a 1:1 population of with representative gametophyte pairs of Macrocystis female and male gametophytes results, which upon maturity pyrifera from seven geographically separated localities in form oogonia and antheridia, producing eggs and spermato- South Chile. They compared the growth performance of all zoids, respectively. Under favourable conditions, zygotes and 49 hybrid constructs starting from zygotes up to the age of young sporophytes appear a few weeks later. 10 weeks and found the following results: As a first step towards improved Macrocystis mariculture – Sporophyte batches produced by intra-population (control) management, Westermeier et al. (2006) introduced a new matings exhibited statistically significant differences in laboratory-based culture technique. They started from field- their growth potentials on relatively low level, which can collected meiospores and isolated individual gametophytes be interpreted as a manifestation of inbreeding depression. with favourable properties: good vegetative growth combined – In contrast, sporophyte batches from several outcrossing with maximum fecundity. Such clonal gametophyte cultures combinations showed significantly higher growth rates can be maintained and propagated vegetatively in unlimited than those of their parents. manner. Gametogenesis can then be initiated at any time by mixing aliquots of both sexes and manipulation of culture These results suggested that Macrocystis sporophytes conditions. This method allows the production of repeated resulting from outbreeding crosses exhibit heterosis, or hybrid batches of up to 105 juvenile sporophytes. Floating freely vigour, which is defined as “the superiority of the offspring of under continuous aeration in flasks, cylinders and tanks, they a cross between two stocks to the better of the parents” (Paul continue to grow and are ready to be explanted to the sea at a 1992).Heterosisisanimportantprincipleusedforcrop length of 8 cm. With this scheme, Westermeier et al. (2006) improvement in terrestrial plants and animal breeding. It is reached 80 kg of Macrocystis biomass m 1 of rope within evident that laboratory-based seedling production in Macro 12 months from the start. This compares favourably with cystis now offers the chance to apply heterosis breeding to a 14.4 kg within 8 months (Gutierrez et al. 2006) and 22 kg marine crop system with commercial significance. within 5 months (Macchiavello et al. 2010) for traditional We report here our attempts to verify the potential of mariculture starting from natural spores seeded on ropes. heterosis breeding for Macrocystis by following favourable In order to further improve the potential of laboratory- heterozygotic sporophyte constructs through their full grown, genetically homogeneous sporophyte seedling commercial mariculture cycles. batches, some details of reproduction biology in Macro cystis must be considered. Graham (2003) and Raimondi et al. (2004) studied natural Macrocystis beds in California. Materials and methods Meiospores have a limited swimming capacity and settle down near their origin to form gametophytes. Survival of Table 1 lists the cultivars, origins and collection dates for the population requires a high density of gametophytes, the Macrocystis gametophyte clones used in this study. It since the distance for successful interaction between egg contains a selection of those M. pyrifera clones originating and sperm does not exceed the range of millimetres. from South Chile (p2 to p7) that were recognized as parents Raimondi et al. (2004) estimated that the area within which of especially successful heterosis crosses in our previous a given sporophyte can reliably produce new progeny only study (Westermeier et al. 2010). For comparison, we added reaches out a few metres from its basal holdfast. This abatchofM. pyrifera sporophytes produced by a mixture of spatial limitation strongly enhances inbreeding, i.e. zygote gametophytes from a natural spore suspension originating formation by gametophytes originating from the same from locality p3. Furthermore, we included one pair of M. parent individual. In consequence, inbreeding-mediated integrifolia parents from North Chile (i8). 821 Table 1 Collection sites, dates and cultivar designations for Locality/cultivar