EFFECT OF HIGH IRRADIANCE ON RECRUITMENT OF THE GIANT KELP MACROCYSTIS (PHAEOPHYTA) IN SHALLOW WATER1

Laboratory and field experiments were done hi Still-water Cove, Carmel Bay, California, and Monterey Harbor, California, to determine the effect of photosynthetically active radiation (PAR) on the shallow (upper) limit of giant kelp, Macrocystis pyrifera (L.) C. Agardh. At shallow depths, M. pyrifera did not recruit or grow to macroscopic size from gametophytes or embryonic sporophytes transplanted to vertical buoy lines; sharp decreases in PAR with depth coincided with observed recruitment and sporophyte distributions. Shade manipulations indicated that settlement of M. pyrifera zoospores was decreased, but not prohibited, by high PAR. Postsettlement stages (gametophytes and embryonic sporophytes), however, survived only under shade. These results suggest that high PAR can inhibit the recruitment of M. pyrifera to shallow water by killing its postsettlement stages; whether or not ultraviolet (UV) radiation also inhibits recruitment was not tested. In either case, however, it appears that high irradiance (PAR and/or UV) regulates the shallow limit of M. pyrifera prior to temperature and desiccation stresses inherent to intertidal regions. In an additional experiment, recruitment or growth of transplanted gametophytes or embryonic sporophytes of Macrocystis integrifolia Bory also did not occur at shallow depths, suggesting that this shallow water species accesses high irradiance regions via a method other than sexual reproduction.     

INTERGENERIC HYBRIDIZATION AMONG FIVE GENERA OF THE FAMILY LESSONIACEAE (PHAEOPHYCEAE) AND EVIDENCE FOR POLYPLOIDY IN A FERTILE PELAGOPHYCUS×MACROCYSTIS HYBRID

Hybridization was attempted by combining gametophytes between intergeneric pairs among the following taxa in the Lessoniaceae: Macrocystis pyrifera ( L.) C. Agardh , M. integrifolia Bory, M. angustifolia Bory , Pelagophycus porra ( Leman) Setch ., Nereocystis luetkeana ( Mert.) Post & Rupr ., Dictyoneurum californicum Rupr ., and Dictyoneuropsis reticulata ( Saud.) Smith. Hybrid sporophytes were produced in some combinations involving Macrocystis × Pelagophycus and Macrocystis × Dictyoneurum, and in all combinations of Dictyoneuropsis × Dictyoneurum. This is the first report of intergeneric hybrids involving Dictyoneurum. Gametophytes of P. porra had 16–24 chromosomes. Gametophytes from a fertile Macrocystis-Pelagophycus hybrid were crossed with Macrocystis and Pelagophycus gametophytes. Hybrid male gametophytes and Pelagophycus female gametophytes produced sporophyte progeny, but hybrid males with Macrocystis females did not. A single hybrid female gametophyte did not produce gametophytes in combination with hybrid males , Pelagophycus males or Macrocystis males. The hybrid gametophytes had approximately 30 chromosomes. It is hypothesized that the hybrid is an alloploid, containing a complete set of Macrocystis and Pelagophycus chromosomes, which may have allowed meiosis and sporogenesis to proceed normally in the hybrid sporophyte found in the sea. Thus, reproductive isolating mechanisms appear to operate at both pre- and postzygotic stages, and both can be overcome in intergeneric hybrids .     

PHENOTYPIC PLASTICITY RECONCILES INCONGRUOUS MOLECULAR AND MORPHOLOGICAL TAXONOMIES: THE GIANT KELP,MACROCYSTIS (LAMINARIALES,PHAEOPHYCEAE), IS A MONOSPECIFIC GENUS1

The giant kelp genus Macrocystis C. Agardh (Laminariales, Phaeophyceae) is one of the world’s most ecologically and economically important seaweed taxa, yet its taxonomy remains uncertain. Although the genus currently contains four accepted species based on variable holdfast and blade morphology [M. pyrifera (L.) C. Agardh, M. integrifolia Bory, M. angustifolia Bory, and M. laevis C. H. Hay], numerous recent studies on Macrocystis interfertility, genetic relatedness, and morphological plasticity all suggest that the genus is monospecific. We reviewed this evidence and present an explanation for the extreme phenotypic plasticity that results in morphological variability within Macrocystis, driven by the effects of environmental factors on early development of macroscopic sporophytes. We propose that the genus be collapsed back to a single species, with nomenclatural priority given to M. pyrifera.     

Effects of chelated iron on oogenesis and vegetative growth of kelp gametophytes (Phaeophyceae)

The supply of iron has been reported to affect gametogenesis in the gametophytes of some species of kelps (order Laminariales). Spores of the kelps Alaria marginata Postels & Ruprecht, Dictyoneurum californicum Ruprecht, Egregia menziesii (Turner) Areschoug, Laminaria setchellii Silva, and Macrocystis pyrifera (L.) C. Agardh were cultured in enriched seawater with and without added chelated iron (Fe‐ethylenediaminetetraacetate) to determine the effects of iron on oogenesis. All species showed a decrease in oogenesis without added Fe‐ethylenediaminetetraacetic acid (EDTA). Gametophytes of E. menziesii showed predominant gametogenesis with or without supplied iron, resulting in all cells being converted to gametes so that vegetative growth did not continue. Vegetative gametophytes were obtained in the other species used. Gametophytes of M. pyrifera did not show any oogenesis without added Fe‐EDTA, while those of L. setchellii, A. marginata and D. californicum were intermediate in their response, showing some gametogenesis without added Fe‐EDTA. When Fe‐EDTA supply was delayed by 6, 13 and 20 days with spores of M. pyrifera, the gametophytes produced fewer eggs, with a greater decrease as the delay grew longer. A range of Fe‐EDTA concentrations was investigated using isolated female gametophytes of two strains of M. pyrifera and one of Macrocystis integrifolia Bory. None of these three strains produced gametes without the addition of Fe‐EDTA. Gametophytes of M. integrifolia required the least amount of added Fe‐EDTA to achieve gametogenesis while gametophytes of M. pyrifera required higher amounts, with the two strains showing somewhat different responses. Iron nutrition appears to be an essential factor for gametogenesis in several species of kelps.     

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.     

ROLE OF NUTRIENT FLUCTUATIONS AND DELAYED DEVELOPMENT IN GAMETOPHYTE REPRODUCTION BY MACROCYSTIS PYRIFERA (PHAEOPHYCEAE) IN SOUTHERN CALIFORNIA1

Organisms occurring in environments subject to severe disturbance and/or periods of poor environmental quality that result in severe adult mortality can survive these periods by relying on alternate life stages that delay their development in a resistant state until conditions improve. In the northeast Pacific, the forest‐forming giant kelp Macrocystis pyrifera (L.) C. Agardh periodically experiences widespread adult mortality during extended periods of extremely low nutrients and high temperatures, such as those associated with El Niño. Recovery following these periods is hypothesized to occur from microscopic life stages that delay their development until the return of favorable conditions. In the laboratory, we experimentally examined the environmental conditions responsible for regulating delayed development of the microscopic stages of M. pyrifera from Southern California, USA. Nutrients controlled the delay and resumption of gametophyte growth and reproduction, perhaps linked to the large fluctuations in nutrients occurring seasonally and interannually in this region. Although growth of gametophytes proceeded in the virtual absence of nitrate, both nitrate and other trace nutrients were necessary for gametogenesis. Upon exposure to elevated nutrients, delayed gametophytes produced sporophytes more quickly (5–20 d) and at smaller sizes (10–200 μm) than gametophytes that had never been delayed (18–80 d, 80–400 μm, respectively), reducing negative density‐dependent effects. This finding demonstrates that delayed gametophytes of M. pyrifera rapidly utilize increased resources to consistently produce sporophytes. Further work is needed to assess their potential role in population recovery following periods of poor environmental quality.     

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.     

A MULTISPECIES LABORATORY ASSESSMENT OF RAPID SPOROPHYTE RECRUITMENT FROM DELAYED KELP GAMETOPHYTES1

Recent work suggests that the ability to delay reproduction as resistant haploid gametophytes may be important for seaweeds that experience unpredictable disturbances or seasonal periods of poor conditions that result in adult sporophyte absence. Further, delayed gametophytes of some kelp species (order Laminariales) may produce sporophytes more rapidly than if they had never experienced a delay, conferring a competitive advantage when conditions improve or after disturbance events. Here, it was determined that the gametophytes of the canopy‐forming kelp Macrocystis pyrifera (L.) C. Agardh could delay reproduction in a one‐ to two‐cell state (<50 μm) for at least 7 months when grown under nutrient‐limiting conditions. These stages retained reproductive viability and produced sporophytes within 5 d once nutrients were increased. This finding suggests that gametophytes could potentially promote recovery of M. pyrifera populations after extended periods of sporophyte absence. In addition, the time required for sporophyte production between gametophytes of the four most conspicuous kelp species in Southern California that had delayed reproduction and gametophytes that had not was compared. For these four kelp species, a delay of at least 30 d conferred a 40%–76% reduction in the time required for sporophyte production once nutrients were received. Fecundity did not decrease with delay duration, suggesting there is no apparent cost of delayed development for kelps as has been observed in other organisms. Thus, delayed development may be a viable strategy for surviving and initially dominating in environments with variable quality.     

AN EVALUATION OF METHODS USED TO ASSESS INTERGENERIC HYBRIDIZATION IN KELP USING PACIFIC LAMINARIALES (PHAEOPHYCEAE)1

Kelp intergeneric laminarialean hybridizations and hybridization protocol were assessed using seven northeast Pacific kelp species: Alaria marginata Postels and Ruprecht, Costaria costata (C. A. Agardh) Saunders, Eisenia arborea Areschoug, Laminaria saccharina (L) Lamouroux, Lessoniopsis littoralis (Tilden) Reinke, Macrocystis integrifolia Bory, and Nereocystis leutkeana (Mertens) Postels and Ruprecht. Survival and development of sporophyte morphologies derived from selfings, separate males and females, and reciprocal crosses were evaluated over 30 weeks of cultivation. All cultures were initiated from cloned gametophytes. Two closely related species, Laminaria angustata Kjellman and L. japonica Areschoug, demonstrated the efficacy of long‐term (up to 30 years) cloned gametophytes in hybridization studies. Sporophyte morphologies appeared in 34%–69% of control and hybridization trials, and 6%–16% of all trials produced sporophytes in control and hybridization conditions that persisted through 30 weeks of cultivation. Sporophytes in control and hybridization conditions could appear normal or abnormal. Usually, the morphology of sporophytes in hybridizations and female controls resembled the female parent, whereas the sporophytes in male controls often had an abbreviated morphology, lacking definitive generic features. Species‐specific rDNA internal transcribed spacer molecular primers were used to determine the parentage of five putative hybrids. Only the L. japonica♀/L. angustata♂ hybrid bore both parental genomes. That negative controls could produce persistent and normal‐appearing sporophytes negates their value and emphasizes the importance of molecular confirmation in hybridization studies. These findings were applied to critique the only known wild intergeneric hybrid, Pelagophycus/Macrocystis.     

Compensatory growth of the kelp Macrocystis integrifolia (Phaeophyceae, Laminariales) against grazing of Peramphithoe femorata (Amphipoda, Ampithoidae) in northern-central Chile

Compensation of tissue loss has been considered an alternative strategy for seaweeds that have no or only minor chemical or structural defense against herbivory. Compensatory responses are facilitated by resource transfer among different tissues and have been suggested for large kelps. Macrocystis integrifolia (Bory) is a common kelp species from northern-central Chile, which is characterized by high growth rates and the absence of lipophilic chemical defenses against herbivore grazing. Herein, we used the giant kelp M. integrifolia to test for compensatory growth in response to grazing by the nest-dwelling amphipod Peramphithoe femorata (Krøyer). Amphipods were allowed to graze inside nests on subapical blades of M. integrifolia sporophytes for 14 days. We measured growth and chemical composition (C, N, laminaran and mannitol) of apical and subapical blades of grazed and ungrazed (control) sporophytes. Our results revealed the capability of M. integrifolia to maintain elongation rates in grazed subapical blades, which were similar to those of subapical blades from ungrazed sporophytes. Apical blades grew slower in grazed than in ungrazed sporophytes indicating a trade-off between apical and subapical blades when herbivores were present. Thus, compensation occurs in blades directly attacked by grazers and is probably mediated by vertical resource allocation within sporophytes to subapical blades, a suggestion supported by the fact that stipe internodes in these regions grew more on grazed sporophytes. In general, our study indicates that M. integrifolia exhibits compensatory growth against the herbivore amphipod P. femorata, and we suggest that this could be an important strategy of large kelp species to tolerate moderate grazing intensities.     

Influence of sedimentation in the absence of macrograzers on recruitment of an annual population of Macrocystis pyrifera in Metri Bay,Chile

The effects of sedimentation and substrate orientation on algal and sessile invertebrate assemblages were tested on an annual population of Macrocystis pyrifera in Metri Bay, southern Chile. In the laboratory, M. pyrifera zoospores were seeded on Crepipatella fecunda shells, the primary substrate for M. pyrifera in this system. The seeded shells were deployed at Metri Bay inside cages and were orientated vertically and horizontally under two sedimentation regimes (bottom and suspended). Due to differences in grazer accessibility and the species present between the sedimentation treatments, grazers (>1 cm) were excluded. We followed sporophyte development of M. pyrifera and the natural recruitment of other algal and invertebrate species. Sedimentation rates were significantly higher in the cages attached to the bottom compared to suspended cages (P < 0.001). In total M. pyrifera and three additional algal genera were detected and all algal recruits showed significantly greater recruitment on the horizontally orientated substrate compared to the vertical substrate. Macrocystis pyrifera sporophytes were present only on the horizontal, suspended (less sedimentation) treatment. In contrast, Ulva and Ectocarpus spp. also occurred in the horizontal, high sediment treatment. Invertebrate recruitment (amphipods, barnacles and spirorbids) dominated the vertically oriented shells regardless of sedimentation. Results indicate that high sedimentation negatively affected the development of M. pyrifera sporophytes while other opportunistic species were able to recruit under these conditions.     

EFFECTS OF LIGHT AND TEMPERATURE ON NET PHOTOSYNTHESIS AND DARK RESPIRATION OF GAMETOPHYTES AND EMBRYONIC SPOROPHYTES OF MACROCYSTIS PYRIFERA1

The rates of net photosynthesis as a function of irradiance and temperature were determined for gametophytes and embryonic sporophytes of the kelp, Macrocystis pyrifera (L.) C. Ag. Gametophytes exhibited higher net photosynthetic rates based on oxygen and pH measurements than their derived embryonic sporophytes, but reached light saturation at comparable irradiance levels. The net photosynthesis of gametophytes reached a maximum of 66.4 mg O₂ g dry wt^?1 h^?1 (86.5 mg CO₂ g dry wt^?1 h^?1), a value approximately seven times the rate reported previously for the adult sporophyte blades. Gametophytes were light saturated at 70 μE m^?2 s^?1 and exhibited a significant decline in photosynthetic performance at irradiances 140 μE m^?1 s^?1. Embryonic sporophytes revealed a maximum photosynthetic capacity of 20.6 mg O₂ g dry wt^?1 h^?1 (25.3 mg CO₂ g dry wt^?1 h^?1), a rate about twice that reported for adult sporophyte blades. Embryonic sporophytes also became light saturated at 70 μE m^?2 s^?1, but unlike their parental gametophytes, failed to exhibit lesser photosynthetic rates at the highest irradiance levels studied; light compensation occurred at 2.8 μE m^?2 s^?1. Light-saturated net photosynthetic rates of gametophytes and embryonic sporophytes varied significantly with temperature. Gametophytes exhibited maximal photosynthesis at 15° to 20° C, whereas embryonic sporophytes maintained comparable rates between 10° and 20° C. Both gametophytes and embryonic sporophytes declined in photosynthetic capacity at 30° C. Dark respiration of gametophytes was uniform from 10° to 25° C, but increased six-fold at 30° C; the rates for embryonic sporophytes were comparable over the entire range of temperatures examined. The broader light and temperature tolerances of the embryonic sporophytes suggest that this stage in the life history of M. pyrifera is well suited for the subtidal benthic environment and for the conditions in the upper levels of the water column.     

PURIFICATION OF SPECIES SPECIFIC ANTIBODIES TO CARBOHYDRATE COMPONENTS OF MACROCYSTIS PYRIFERA (PHAEOPHYTA)1

Polyclonal rabbit antibodies to cell wall components were produced against gametophytes of the giant kelp Macrocystis pyrifera (Linnaeus) C. Agardh. These antibodies were found to react with carbohydrates extracted from M. pyrifera and Pterygophora californica Ruprecht by carbohydrate based enzyme immunoassay (EIA). The antibodies reacted with carbohydrates from both species. After affinity purification on a column with M. pyrifera carbohydrate coupled to AH-Sepharose, the eluted antibody was specific for M. pyrifera carbohydrate with little cross reactivity to P. californica carbohydrate in the EIA test. In experiments carried out to characterize the antigenic specificity of unfractionated antibody using commercially prepared carbohydrates in the EIA, the antibodies were shown to react primarily with fucoidan and to a lesser degree, alginate. The unfractionated antibody was also shown to bind to proteins from both M. pyrifera and P. californica. These results indicate that species specific carbohydrate determinants may be present in the kelp cell wall.     

Natural variation of Macrocystis pyrifera gametophyte germplasm culture microbiomes and applications for improving yield in offshore farms

With national interest in seaweed-based biofuels as a sustainable alternative to fossil fuels, there is a need for tools that produce high-yield seaweed cultivars and increase the efficiency of offshore farms. Several agricultural studies have demonstrated that the application of microbial inoculants at an early life stage can improve crop yield, and there is an opportunity to use similar techniques in seaweed aquaculture. However, there is a critical knowledge gap regarding host–microbiome associations of macroalgae gametophytes in germplasm cultures. Here, we investigate the microbial community of Macrocystis pyrifera gametophyte germplasm cultures that were used to cultivate an offshore farm in Santa Barbara, California and identify key taxa correlated with increased biomass of mature sporophytes. This work provides a valuable knowledge base for the development of microbial inoculants that produce high-biomass M. pyrifera cultivars to ultimately be used as biofuel feedstocks.     

GENETIC LOAD IN OSMUNDA REGALIS POPULATIONS

Osmunda regalis sporophytes form haploid spores which develop into functionally hermaphroditic gametophytes. The self-fertilization of such gametophytes results in zygotes which are completely homozygous. Spore samples collected from sporophytes in natural populations were used to establish gametophyte cultures. The majority of these gametophytes were unable to form viable embryos when only self-fertilization was possible. Controlled selfing and crossing experiments revealed that the inability of these homozygous embryos to develop normally is attributable to the presence of recessive lethals. To account for this genetic load, an hypothesis is proposed integrating the morphology and ecology of the gametophyte generation with the polyploid genetic system of the sporophyte generation.     

ROLE OF SETTLEMENT DENSITY ON GAMETOPHYTE GROWTH AND REPRODUCTION IN THE KELPS PTERYGOPHORA CALIFORNICA AND MACROCYSTIS PYRIFERA (PHAEOPHYCEAE)1

Laboratory studies were used to examine how variation in the density of spore settlement influences gametophyte growth, reproduction, and subsequent sporophyte production in the kelps Pterygophora californica Ruprecht and Macrocystis pyrifera (L.) C. Ag. In still (non-aerated) cultures, egg maturation in both species was delayed when spores were seeded at densities 300 · mm^?2. Although the density at which this inhibition was first observed was similar for both species, the age at which their eggs matured was not. P. californica females reached sexual maturity an average of 4 days (or ～ 30%) sooner than did M, pyrifera. As observed previously in field experiments, per capita sporophyte production was negatively density dependent for both species when seeded at spore densities of 10 · mm^?2. Total sporophyte production (i.e. number · cm^?2) for both species, however, was greatest at intermediate densities of spore settlement (～ 50 spores · mm^?2). In contrast, total sporophyte production by P. californica steadily increased with increasing spore density in aerated cultures; highest sporophyte density was observed on slides seeded at a density of 1000 spores · mm^?2. Preliminary experiments with P. californica involving manipulation of aeration and nutrients indicate that inhibition of gametophyte growth and reproduction at higher densities of spore settlement in non-aerated cultures was probably caused by nutrient limitation.     

Nuclear DNA level and life cycle of kelps: Evidence for sex‐specific polyteny in Macrocystis (Laminariales,Phaeophyceae)

Giant kelp, Macrocystis pyrifera (Linnaeus) C. Agardh, is the subject of intense breeding studies for marine biomass production and conservation of natural resources. In this context, six gametophyte pairs and a sporophyte offspring of Macrocystis from South America were analyzed by flow cytometry. Minimum relative DNA content per cell (1C) was found in five males. Unexpectedly, nuclei of all female gametophytes contained approximately double the DNA content (2C) of males; the male gametophyte from one locality also contained 2C, likely a spontaneous natural diploid variant. The results illustrate a sex‐specific difference in nuclear DNA content among Macrocystis gametophytes, with the chromosomes of the females in a polytenic condition. This correlates with significantly larger cell sizes in female gametophytes compared to males and resource allocation in oogamous reproduction. The results provide key information for the interpretation of DNA measurements in kelp life cycle stages and prompt further research on the regulation of the cell cycle, metabolic activity, sex determination, and sporophyte development.     

Sporophyte formation in experimentally-induced unisexual female and bisexual gametophytes ofLygodium japonicum

Unisexual female and male and bisexual gametophytes were experimentally induced inLygodium japonicum. A single bisexual gametophyte was isolated in a dish and a female gametophyte was paired with a male one to allow intragametophytic selfing and intergametophytic mating, respectively. About 30% of the females formed sporophytes but no bisexual gametophytes formed them.     

PHOTOSYNTHETIC PHYSIOLOGY AND CHEMICAL COMPOSITION OF SPORES OF THE KELPS MACROCYSTIS PYRIFERA,NEREOCYSTIS LUETKEANA,LAMINARIA FARLOWII,AND PTERYGOPHORA CALIFORNICA (PHAEOPHYCEAE)1

Recently released spores of the kelps Macrocystis pyrifera (L.) C. Ag., Nereocystis luetkeana (Mert.) Post. and Rupr., Laminaria farlowii Setch., and Pterygophora californica Rupr. had different levels of net photosynthesis. Spore-specific photosynthesis–irradiance relationships were similar in many respects for M. pyrifera, N. luetkeana, and L. farlowii spores. All three species had low rates of net light-saturated photosynthesis. In contrast, spores of P. californica had higher photosynthetic potential and overall net photosynthesis than the other three species. On a cell carbon basis, however, photosynthetic rates in N. luetkeana spores were similar to those of P. californica spores and higher than those of M. pyrifera spores. Chlorophyll a content of spores varied 10-fold among species. The rank order of significant differences in chlorophyll a content was P. californica > L. farlowii > N. luetkeana > M. pyrifera. As a result, chlorophyll-specific measurements suggest M. pyrifera and N. luetkeana spores had much higher quantum efficiency and photosynthetic potential than either P. californica or L. farlowii spores. Maternal carbon and nitrogen investment significantly differed in spores of M. pyrifera, N. luetkeana, and P. californica with P. californica > M. pyrifera > N. luetkeana. Carbon content in spores of each of these three species increased by about 30% during 12 h of saturating irradiance. We suggest that the photosynthetic capabilities of and maternal investment in spores may be related to the spore as a unit of dispersal, to the reproductive ecology of the parental sporophytic stages, and to the growth and physiology of the germling gametophyte stages.