Diego Aedo

CONVERGENT BIOLOGICAL PROCESSES IN COALESCING RHODOPHYTA

Sporeling coalescence in Gracilaria chilensis Bird, McLachlan et Oliveira produces genetically polymorphic, chimeric individuals. If this is common in red algae, it may have significant biological consequences. In this study, we evaluate the hypotheses that coalescence is widespread among the Rhodophyta and that specific and convergent morphological and ecological responses characterize this as a unique growth style among marine algae. A literature survey on coalescence was undertaken to assess the distribution of this condition in the Florideophycidae. Sixty‐two (54.9%) of 113 species considered germinated to form a disk. Subsequent development in 37 of these species showed crust formation and coalescence during development with other crusts in 31 species (84%). Coalescing red algae were members of the orders Ahnfeltiales, Corallinales, Gigartinales, Gracilariales, Halymeniales, Palmariales, and Rhodymeniales. Ultrastructural studies in species of Ahnfeltiopsis, Chondrus, Gracilaria, Mazzaella, and Sarcothalia suggested a common pattern of early development. Newly released, naked spores may fuse into a single cell, as they do in Chondrus canaliculatus, or they may develop individual cell walls that later are surrounded by a thickened common wall. Ultrastructural studies demonstrated two kinds of immediate development after the first mitotic division: direct development by symmetric divisions resulting in discoid sporelings or an indirect asymmetric arrangement of divisions before a diskoid sporeling was formed. Germination in coalescing species is a linear function of the initial spore density, whereas in noncoalescing species maximum germination occurs at intermediate densities. In the field, coalescing species may recruit either from solitary or aggregated spores. However, survival is significantly higher for plantlets grown from a larger number of coalescing spores. Total number of erect axes formed by the coalesced mass is a logarithmic function of the initial number of spores. Thus, germlings grown from a larger number of coalescing spores exhibited a larger photosynthetic canopy than do plantlets grown from a few spores. Juveniles and mature clumps grown from a coalescing mass may exhibit size inequalities among erect axes, with the larger axes located toward the center of the clump. These larger axes mature first or, in some cases, are the only to produce spores. The widespread occurrence of coalescence inroughly half the number of orders of the Florideophycidae, the similarity of the coalescence process, and the finding of various adaptive traits associated with coalescence characterizes this as a unique growth style, splitting the diversity of species now included in the Florideophycidae into two major groups: coalescing and noncoalescing Rhodophyta.

SPORELING COALESCENCE AND INTRACLONAL VARIATION IN GRACILARIA CHILENSIS (GRACILARIALES, RHODOPHYTA)

This study evaluates the hypothesis that spore coalescence may cause intraclonal variation. Spore coalescence might allow the occurrence of unitary thalli that in fact correspond to genetically different, coalesced individuals. Plant portions simultaneously derived from these chimeric individuals may exhibit dissimilar growth responses even when incubated under similar abiotic conditions. Testing of the hypothesis included various approaches. Transmission electron microscopy observations of early stages of sporeling coalescence indicated that polysporic plantlets were formed by groups of spores and their derivatives. Even though adjacent cells in two different groups may fuse, these groups maintained an independent capacity to grow and form uprights. Laboratory‐grown plantlets showed a significant correlation between the initial number of spores and the total number of erect axes differentiated from the sporeling. Construction and growth of bicolor individuals indicated the chimeric nature of the coalesced individuals. Coalesced, bicolor holdfasts had green and red cells, which subsequently produced green and red uprights, respectively. Individuals fronds were also chimeric, as indicated by the production of green and red branchlets from single, red uprights. The existence of mixed tissues was further substantiated by random amplified polymorphic DNA analysis. The banding pattern produced by branchlets of a unisporic thallus was consistently monomorphic, whereas the patterns produced by the polysporic thallus were polymorphic. Growth rates of polysporic thalli had larger data dispersal and variation coefficients than oligosporic or monosporic thalli. Therefore, all results support the original hypothesis and suggest that coalescence might be ecologically more important than previously thought.

Causes and implications of intra-clonal variation in Gracilaria chilensis (Rhodophyta)

Strain selection studies in Gracilaria chilensis detected significant levels of intra-clonal variation. These findings motivated more detailed studies on the causes and implications of intra-clonal variation in these and other red algal species. Our results indicate that intra-clonal variation is common among replicated units (e.g.: carpospores and ramets) of several red algal species and suggest that a larger data base probably will show the occurrence of various kinds of intra-clonal changes, differing in frequency of occurrence and magnitude of phenotypic expression. It is likely also that different species would exhibit different amounts of variation. Four types of factors may cause intra-clonal variation: (1) physiological or developmental differences among ramets, (2) localized pathogen infections, (3) several kinds of genetic changes, and (4) sporeling coalescence. Intra-clonal variation among ramets: (1) increases the possibility of genet survival, (2) explains the origin of morphological and physiological differences among ramets of a given genet, (3) explains the large population variation found in many clonal species and (4) suggests that strain selection of some economically important seaweeds should be thought of as a fairly continuous process due to the instability of some of these clones.

Banks of microscopic forms and survival to darkness of propagules and microscopic stages of macroalgae

Estudios previos han encontrado que el numero de especies formando un banco de formas microscopicas en pozas de mareas de Chile central incluyo solo la mitad del numero de especies presentes en la vegetacion macroscopica en las cercanias de las pozas intermareales. Una primera condicion para sobrevivir en estos bancos radica en la capacidad de las formas microscopicas para tolerar oscuridad total o baja iluminacion por periodos prolongados. Para evaluar dicha capacidad, los propagulos de 17 especies de algas verdes, pardas y rojas, presentes y ausentes del banco de formas microscopicas fueron incubadas a distintas combinaciones de intensidad luminosa y fotoperiodo. Propagulos del 47 % de las especies evaluadas (ocho especies) germinaron en oscuridad mientras que los propagulos de las otras nueve especies requirieron valores muy bajos de intensidad luminosa (2-10 µmol m-2 s-1) para germinar. En una mayoria de las especies, las formas microscopicas mostraron una mayor tolerancia a la oscuridad que los propagulos. Algunos sobrevivieron en la oscuridad por sobre un ano y una especie (Gelidium lingulatum) pudo sobrevivir en oscuridad absoluta por 500 dias. La habilidad para sobrevivir en oscuridad total no se relaciona con presencia de la especie en los bancos de formas microscopicas, con grupos filogeneticos o con historias de vida especificas, con tamano de propagulo, morfologia de la forma microscopica o estatus sucesional (especies fugitivas versus sucesionales tardias). Por lo tanto, tolerancia a la oscuridad aparece como un patron comun a propagulos y formas microscopicas de una mayoria de algas bentonicas. Los patrones de crecimiento exhibidos por las formas microscopicas de Lessonia nigrescens, Chaetomorpha firma y Glossophora kunthii sugiere que el efecto de altas intensidades luminosas sobre estos reclutas podria determinar los limites superiores de distribucion vertical de estas especies

Cultivation of Gigartina skottsbergii (Gigartinales, Rhodophyta): Recent advances and challenges for the future

This study integrates landings statistics and biological studies of the red algaGigartina skottsbergii Setchell & Gardner. The analysis of the landings and carrageenan production in Chile suggeststhat this resource will suffer a strong harvesting pressure during the nextyears. Biological results on sporulation, germination, sporeling growth and survivorship in laboratory,indoor tanks and field conditions, indicated that cultivation of this species istechnically feasible, as spores can be seeded on ropes and other substrata. Vegetative propagation of this species through tissue fragmentationis also possible. Vegetative fragments of this carrageenophyte have 20 to30% higher growth rates than whole fronds in suspended culture systems. Protoplast production can be also explored for bypassing restrictions inspore availability. Major advantages that encourage the cultivation of G. skottsbergii include its gel quantity and quality, its pathogen-freecondition, a high reproduction potential and its regeneration capacity. Onthe other hand, the major constraints are related to its relatively slowgrowth as compared to other carrageenophytes, limited availability ofspores and high mortality during juvenile stages.

Group Recruitment and Early Survival of Mazzaella Laminarioides

Several phycocolloid-producing Rhodophyta of significant economic importance are coalescing species, able to fuse with conspecifics during recruitment, reach larger sizes and increase their survival. In these species spores are needed to start cultivation (e.g. Gigartina, Mazzaella) or to increase the seed stocks, to renew senescent clones or to enlarge the base of genetic variation of vegetatively propagated species (e.g. Chondrus, Gracilaria, Eucheuma). This study uses Mazzaella laminarioides to evaluate some key features that influence recruitment success. Field measurements indicate that in any recruitment event a variable amount of the spores reaching a given place may form groups of 2 to over 100 coalescing spores, while field experiments support the idea that early recruitment success is a function of the number of coalescing spores forming the individual, as multisporic, coalescing recruits have higher survival rates than sporelings formed by one or a few spores. Therefore, group recruitment (spores settling and recruiting in close spatial proximity) appears as a prerequisite for sporeling coalescence and early recruitment success. In turn, laboratory experiments suggest that the frequency of group recruitment and coalescence increases with increasing spore abundance and with slight Ca++ additions to the culture medium. These last two factors could be handled by farmers to improve the success of spore inoculations of coalescing species.