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.     

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.     

PHYSIOLOGICAL PERFORMANCE OF FLOATING GIANT KELP MACROCYSTIS PYRIFERA (PHAEOPHYCEAE): LATITUDINAL VARIABILITY IN THE EFFECTS OF TEMPERATURE AND GRAZING1

Rafts of Macrocystis pyrifera (L.) C. Agardh can act as an important dispersal vehicle for a multitude of organisms, but this mechanism requires prolonged persistence of floating kelps at the sea surface. When detached, kelps become transferred into higher temperature and irradiance regimes at the sea surface, which may negatively affect kelp physiology and thus their ability to persist for long periods after detachment. To examine the effect of water temperature and herbivory on the photosynthetic performance, pigment composition, carbonic anhydrase (CA) activity, and the nitrogen (N) and carbon (C) content of floating M. pyrifera, experiments were conducted at three sites (20° S, 30° S, 40° S) along the Chilean Pacific coast. Sporophytes of M. pyrifera were maintained at three different temperatures (ambient, ambient ? 4°C, ambient + 4°C) and in presence or absence of the amphipod Peramphithoe femorata for 14 d. CA activity decreased at 20° S and 30° S, where water temperatures and irradiances were highest. At both sites, pigment contents were substantially lower in the experimental algae than in the initial algae, an effect that was enhanced by grazers. Floating kelps at 20° S could not withstand water temperatures >24°C and sank at day 5 of experimentation. Maximal quantum yield decreased at 20° S and 30° S but remained high at 40° S. It is concluded that environmental stress is low for kelps floating under moderate temperature and irradiance conditions (i.e., at 40° S), ensuring their physiological integrity at the sea surface and, consequently, a high dispersal potential for associated biota.     

EFFECTS OF THE ENCRUSTING BRYOZOAN,MEMBRANIPORA MEMBRANACEA,ON THE LOSS OF BLADES AND FRONDS BY THE GIANT KELP,MACROCYSTIS PYRIFERA (LAMINARIALES)1

Juvenile sporophytes of the giant kelp, Macrocystis pyrifera (L.) C. A. Agardh, were transplanted from local kelp beds to stations located various distances from the outfall from an electrical generating station that was known to cause an increase in the settlement of fouling organisms. Plants near the outfall became heavily fouled by the encrusting bryozoan, Membranipora membranacea (L.), and lost about one-third of their blades during the course of the experiment. Blade loss was significantly correlated with amount of fouling. To test the hypothesis that fouling causes blade loss, we paired fouled and unfouled plants of about the same age, overall length, and number of fronds and placed them at stations in nearby kelp beds and near the outfall. At the stations in the kelp beds, the fouled plants lost blades more rapidly than the unfouled controls. However, at the station near the outfall the “control” plants quickly became fouled so there was little difference in treatments and there was no significant difference in blade loss. Plants fouled by Membranipora suffered greater blade loss than clean plants probably because fouled blades are fragile and break off easily and because fish bite off chunks of blade while foraging on the attached bryozoans.     

VARIATION IN NITROGEN PHYSIOLOGY AND GROWTH AMONG GEOGRAPHICALLY ISOLATED POPULATIONS OF THE GIANT KELP,MACROCYSTIS PYRIFERA (PHAEOPHYTA)1

Three geographically isolated populations of the giant kelp, Macrocystis pyrifera (L.) C. Ag., were examined for responses to nitrate availability in batch culture experiments using juvenile sporophytes reared from spores in the laboratory. Although maximum rates of nitrate-saturated growth were similar among groups, there were significant quantitative differences in the response to nitrate limitation that can be related to natural patterns of nutrient availability at these sites. Plants from Santa Catalina Island (most oligotrophic) achieved maximum growth rates at ambient nitrate concentrations that were lower than those for plants from Monterey Bay, California (most eutrophic), or Refugio State Beach (near Santa Barbara, California). Tissue nitrogen and amino acid concentrations were highest in plants cultured from Santa Catalina Island populations at all external nitrate concentrations, suggesting that differences in nitrate requirements for growth may reflect the efficiency of nitrate uptake and assimilation at subsaturating nitrate concentrations. Given the different physical environments from which these plants came, the data suggest that geographically isolated populations of M. pyrifera have undergone genetic divergence that can be explained by ecotypic adaptation to unique habitat conditions at these sites.     

TRANSLOCATION OF 11C-PHOTOASSIMILATE IN THE BLADE OF MACROCYSTIS PYRIFERA (PHAEOPHYCEAE)1

Radioactive bicarbonate was pulse fed to blades of Macrocystis pyrifera (L.) C. A. Ag. and the movement of the ¹¹C-labelled photoassimilates was monitored in vivo using an externally mounted array of Geiger-Müller detectors. Results of experiments conducted in August 1982 and February 1983 showed kinetic transport profiles composed of short pulses of ¹¹C (periods of two to three minutes and six to eight minutes) and a mass flow component travelling with a speed of 6–22 cm · h^?1. The pulse-like movement of ¹¹C-photoassimilates, revealed for the first time in a kelp, may be driven by an energy-assisted transport mechanism. Light microscopy revealed a putative symplastic transport pathway from the photo synthetic meristoderm to the medullary sieve cells in the M. pyrifera blade. Of particular importance were the connections between the inner cortical cells and thin-walled medullary sieve cells. Electron microscopy showed sieve plate pore diameters ranging between 35–60 nm in the cortex and ca. 40 nm in the end walls of the thin-walled sieve cells.     

NEUTRAL LIPIDS AS MAJOR STORAGE PRODUCTS IN ZOOSPORES OF THE GIANT KELP MACROCYSTIS PYRIFERA (PHAEOPHYCEAE)1

Neutral lipids, consisting primarily of triacylglycerols, were found to be a major form of carbon reserve in zoospores of the giant kelp Macrocystis pyrifera (L.) C. Ag. The fluorescent stain Nile Red revealed large lipid droplets in the posterior end of the cell, which comprised 20–41% of cellular carbon in newly released spores. Flow cytometric analyses of newly released spores stained with Nile Red revealed considerable variation in the neutral lipid content among spores that was independent of spore size. Lipid droplets were consumed during germination in spores maintained either under constant light or in continual darkness. The availability of light appeared to delay, but did not preclude, lipid use. The rate of lipid use during germination varied considerably among germlings with some cells consuming all of their lipid reserves within 5 h after release. In addition to zoospores, lipid droplets were observed in both male and female gametes. Numerous droplets were observed in eggs, while single lipid droplets were observed in sperm. Neutral lipid droplets were not observed in gametophytes or sporophytes except in developing gametes and spores. Large lipid reserves thus seem to be confined to the microscopic life history stages that presumably have relatively high energy demands. By serving as a supplemental fuel reserve, neutral lipids may be important in extending the effective range of zoospore dispersal.     

PHOTOSYNTHETIC RESPONSE OF THE GIANT KELP MACROCYSTIS PYRIFERA (PHAEOPHYCEAE) TO ULTRAVIOLET RADIATION1

Solar ultraviolet radiation (UVA + UVB) impairs photosynthesis in marine algae. Canopy blades of the giant kelp Macrocystis pyrifera (L.) C. Agardh are exposed to high levels of solar UV in the field. To determine the effects of UV radiation on photosynthesis in the giant kelp and to identify sites of UV damage, O₂ evolution, reaction center organization, light harvesting, and energy transfer efficiency were measured in canopy blades that had been exposed to elevated levels of UV in the laboratory. UV treatment reduced both the light-saturated rate and the light-limited rate of photosynthesis by 50% but produced no significant change in the rate of dark respiration. A significant impairment of photosystem II (PSII) reaction center function was observed, suggesting that PSII is a major site of damage in chromophytes. Reduced quantum efficiency of photosynthesis and loss of energy transfer from light-harvesting pigments (fucoxanthin, chlorophyll a, and chlorophyll c) to PSII indicate that the major light-harvesting complex of M. pyrifera, the fucoxanthin-chlorophyll protein complex (FCPC), was another site of UV damage. These measures provide the first evidence of a direct effect of UV radiation on specific sites in the photosynthetic apparatus of chromophytes and indicate that in situ fluorescence excitation analysis may be a simple means to detect UV stress in algae.     

SPORE SUPPLY AND HABITAT AVAILABILITY AS SOURCES OF RECRUITMENT LIMITATION IN THE GIANT KELP MACROCYSTIS PYRIFERA (PHAEOPHYCEAE)1

The causes of spatial variation in the recruitment of benthic marine algae are frequently misunderstood because of difficulties in distinguishing among the many factors that influence the supply and establishment of microscopic propagules. We used the recently constructed San Clemente Artificial Reef (SCAR) experiment to examine the roles of dispersal distance, size of spore source, and habitat availability as sources of variation in the recruitment of the giant kelp Macrocystis pyrifera (L.) C. Ag., a species whose recruitment has often been considered to be dispersal limited. Sparse colonization on SCAR by adult Macrocystis occurred within 6 months after reef construction via drifters (i.e. individuals from neighboring kelp beds that became dislodged and set adrift). The abundance of drifters on SCAR declined exponentially with distance from the nearest source population (San Mateo), suggesting that San Mateo was the likely source of drifters. Dense recruitment of small Macrocystis sporophytes was observed within 8 months of reef construction. The density of recruits on SCAR showed an initial increase with distance from San Mateo before declining exponentially. Nonetheless, substantial recruitment was observed at the most distant locations on SCAR located 3.5 km from San Mateo. In contrast to drifters, the density of recruits was positively correlated to the bottom cover of artificial reef substrate. Importantly, no correlation was found between the local density or fecundity of drifters and the local density of kelp recruits suggesting that recruitment on SCAR resulted from widespread spore dispersal rather than from the local dispersal of spores from drifters.     

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.     

ARRESTED DEVELOPMENT OF GIANT KELP (MACROCYSTIS PYRIFERA,PHAEOPHYCEAE) EMBRYONIC SPOROPHYTES: A MECHANISM FOR DELAYED RECRUITMENT IN PERENNIAL KELPS?1

Delayed recruitment of microscopic stages in response to cyclical cues is critical to the population dynamics of many annual and seasonally reproducing perennial seaweeds. Microscopic stages may play a similar role in continuously reproducing perennials in which adult sporophytes are subject to episodic mortality, if they can respond directly to the unpredictable onset and relaxation of unfavorable conditions. We experimentally evaluated the potential for temporary reduction in limiting resources (light, nutrients) to directly delay recruitment of giant kelp (Macrocystis pyrifera (L.) C.A. Agardh) gametophytes and embryonic sporophytes. Laboratory cultures were subjected to limiting conditions of light and nutrients for 1 month and then exposed to nonlimiting conditions for 10 days. Gametophytes in all treatments failed to recruit to sporophytes after 2 weeks, suggesting they are not a source of delayed recruitment in giant kelp. Sporophytes in light‐limited treatments, however, survived and grew significantly slower than non–light‐limited controls. When stimulated with light, light‐limited sporophytes grew from 2 to>10 times faster than unstimulated controls depending on nutrient availability. These results suggest that limiting resources can delay recruitment of embryonic giant kelp sporophytes for at least 1 month. Flexible timing of recruitment from embryonic sporophytes may enhance persistence of continuously reproducing perennial species when mac‐ roscopic adults are subject to episodic large‐scale removals.     

STRUCTURE AND DEVELOPMENT OF THE SECRETORY CELLS AND DUCT SYSTEM IN MACROCYSTIS PYRIFERA (L.) C. A. AGARDH1

Sporophytes of Macrocystis pyrifera (L.) C. A. Agardh of various stages of growth were studied by light microscopy to determine the initiation and ontogeny of secretory cells and the accompanying duct system. Secretory cells are initiated by asymmetric, periclinal divisions of meristoderm cells; subsequent mitoses increase the number of secretory cells associated with each duct. Duct formation occurs by schizogeny of anticlinal cell walls adjacent to the site of secretory cell initiation. Differences in distribution and structure of the duct system occur in various parts of the sporophyte. The duct system does not have openings directly to the sporophyte surface. Histochemical techniques showed that the duct contents are mostly sulfated polysaccharides with perhaps some lipid.     

REPRODUCTIVE LONGEVITY OF DRIFTING KELP MACROCYSTIS PYRIFERA (PHAEOPHYCEAE) IN MONTEREY BAY,USA1

Drifting Macrocystis pyrifera (L.) C. Agardh sporophytes have long been viewed as the primary long‐distance dispersal vector; yet, few data exist that support the ability of reproductive viable sporophytes to actually travel the presumed hundreds to thousands of kilometers. This study addressed the reproductive longevity of experimental and naturally occurring M. pyrifera drifters. Temporal variability in sporophyte size and reproduction was estimated for experimental drifting sporophytes that were tethered to surface buoys and compared with attached plants (controls). Reproductive viability was also studied for beach‐cast drifters (BCD), and naturally drifting sporophytes observed during field surveys in Monterey Bay. Detached drifting sporophytes were tracked with radio transmitters to follow drifter trajectories and to measure drifting speed. Experimental drifters (ED) experienced a 74% reduction in frond length after 35 days, a 76% reduction in average frond number after 70 days, and a reduction in average sorus area by 83% after 28 days. Although zoospore production was reduced following detachment, sporophytes remained fertile with high zoospore germination success as long as sori were present (125 days). Zoospore production and germination success for natural and BCD was similar to ED. The average displacement of radio‐tagged drifters was 7.12 km·day^?1, suggesting that a sporophyte adrift for 125 days disperses viable propagules (zoospores) over 890 km (±363). Dispersal of propagules is important for population restoration, distribution, and genetic diversity. Such dispersal distances are long enough to connect potentially all Northern Hemisphere Macrocystis populations across a generational timescale and may facilitate inter‐hemispheric gene flow.     

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.     

PHOSPHORUS AND THE GROWTH OF JUVENILE MACROCYSTIS PYRIFERA (PHAEOPHYTA) SPOROPHYTES1

The effect of phosphate (P_i) supply on growth rate and tissue phosphorus content of juvenile Macrocystis pyrifera (L.) C. Ag. sporophytes was examined. Sporophytes were batch cultured in aquaria with flowing recirculated seawater enriched by 30 μM nitrate. Each aquarium was supplemented with a different seawater P_i concentration, 0, 0.3, 1, 2, 3, and 6 μM. Sporophyte mean specific growth rates declined with time in all cultures presumably due to the normal developmental decrease in the proportion of meristematic tissue of each plant. Growth rate declines were more pronounced in cultures that were nutrient limited. Sporophyte growth was P-limited after two-week exposure to P_i less than 1 μM, corresponding to a tissue P concentration of less than 0.20% dry weight. Plants cultured at 6 μM P_i contained tissue P levels of 0.53% dry weight after three weeks. Luxury consumption and storage of P occurred.     

GRAZING ON GIANT KELP MICROSCOPIC PHASES AND THE RECRUITMENT SUCCESS OF ANNUAL POPULATIONS OF MACROCYSTIS PYRIFERA (LAMINARIALES,PHAEOPHYTA) IN SOUTHERN CHILE1

The giant kelp Macrocystis pyrifera (L.) C. Agardh is widely distributed in the Northern Hemisphere and Southern Hemisphere, yet it exhibits distinct population dynamics at local to regional spatial scales. Giant kelp populations are typically perennial with the potential for year‐round reproduction and recruitment. In southern Chile, however, annual giant kelp populations exist and often persist entirely on secondary substrata (e.g., shells of the slipper limpet Crepipatella fecunda [Gastropoda, Calyptraeidae]) that can cover up to 90% of the rocky bottom. In these populations, the macroscopic sporophyte phase disappears annually during winter and early spring, leaving a 3–4 month period in which a persistent microscopic phase remains to support the subsequent year’s recruitment. We tested the effects of a suite of grazers on the recruitment success of this critical microscopic phase at two sites in southern Chile. Field experiments indicated that the snail Tegula atra negatively impacted M. pyrifera sporophyte recruitment, but that recruitment was highest in the presence of sessile female limpets, C. fecunda. Conversely, small male C. fecunda (biofilm grazers) did not regulate kelp recruitment. Laboratory observations showed that C. fecunda males only grazed on microscopic kelp gametophytes and small (<250 μm) sporophytes, rejecting larger sporophytes, whereas T. atra grazed on all the kelp stages. Recruitment to the C. fecunda treatments far exceeded that to bare rock in the absence of grazers but was not due to the physical presence of C. fecunda shells. We concluded that the key to M. pyrifera recruitment success in southern Chile is its capacity to colonize secondary substrates provided by the slipper limpet C. fecunda.     

IRRADIANCE,DAYLENGTH AND TEMPERATURE EFFECTS ON ZOOSPOROGENESIS IN COLEOCHAETES CUTATA (CHAROPHYCEAE) 1

Using a factorial design, we investigated the effects of 150 different combinations of irradiance, daylength and temperature on zoosporogenesis in Coleochaete scutata. Analysis of variance (ANOVA) revealed that irradiance and daylength did not .significantly influence the response, out that temperature was highly significant. Exposure of thalli to 20°C for one to several days is sufficient to induce zoospore production in C. scutata and several other northern temperature species of Coleochaete. Results of the factorial experiment correlate well with field observations on the seasonal occurrence of asexual reproduction in several Coleochaete species. A technique based on results of this factorial study is described for using zoospores to obtain morphologically normal, unialgal cultures of Coleochaete .sp. It was concluded that the factorial approach to investigation of environmental control of zoosporogenesis can be a powerful tool for understanding natural algal population dynamics, as well as controlling growth and reproduction of algae in the laboratory.     

CRITICAL LEVELS OF LIGHT AND TEMPERATURE REGULATING THE GAMETOGENESIS OF THREE LAMINARIA SPECIES (PHAEOPHYCEAE)1

Gametophytes of three Laminaria species occurring near Helgoland, North Sea, were cultivated 4 wk in a 12:12 LD regime at different temperatures in artificial light fields, and in the sea at different water depths. In the artificial light fields underwater spectral distribution was simulated according to Jerlov water Types 5, 7, 9. Blue light in the simulated light fields amounted to 17, 12 or 4% of total quanta. The rate of vegetative growth did not depend on spectral distribution, was light-saturated at 4–6 W · m^?2, and increased with temperature up to 15 C. L. saccharina (L.) Lamour. exhibited the highest tolerance towards temperature, light and UV. Gametophytes survived 1 wk at 21 C ± 0.1, but not 22 C ± 0.1. Gametophytes of L. hyperborea (Gunn.) Fosl. and L. digitata (Huds.) Lamour. survived 1 wk at 20 C ± 0.1, but not at 21 C ± 0.1. In sunlight, and in the light field of a xenon lamp, 50% of L. saccharina gametophytes were killed by a quantum dose of 50 μEin · cm^?2, and 100% of the plants by 90 μEin · cm^?2. Approximately half of these quantum doses killed the corresponding percent of the other species gametophytes. Appreciably higher quantum doses were survived in visible light, with red being the most detrimental. Fertility depended on a critical quantum dose of blue light which decreased almost exponentially with decreasing temperature. The quantum dose (400–512 nm) required for induction of fertilization of 50% of the female gametophytes (males react similarly) was 90 μEin · cm^?2 at 5 C, 110 μEin · cm^?2 at 10 C, 230 (560 in L. digitata)μEin · cm^?2 at 15 C, and 560 (L. hyperborea) or about 850 (other 2 species) μEin · cm^?2 at 18 C. In the sea the gametophytes survived the dark winter months in the unicellular stage, with almost no vegetative growth of the primary cell, due to lack of light. In early spring the female gametophytes matured in the unicellular, and the males in a few-celled stage at the depth of 2 m, as did the laboratory cultures under conditions inducing maximal fertility.