CRITICAL IRRADIANCE LEVELS AND THE INTERACTIVE EFFECTS OF QUANTUM IRRADIANCE AND DOSE ON GAMETOGENESIS IN THE GIANT KELP,MACROCYSTIS PYRIFERA1

Gametophytes of Macrocystis pyrifera (L.) C. Ag. were cultured under a series of quantum irradiances in three photoperiod regimes. The quantum irradiances in each photoperiod were adjusted to provide equal daily irradiation dosages between photoperiods which allowed a critical examination of the interactions between quantum irradiance and quantum dose in determining gametophyte fertility. The lowest quantum irradiance which stimulated gametogenesis in more than 50% of the female gametophytes was 5 μE·m^?2·s^?1. The saturating irradiance was ca. 10 μE·m^?2·s^?1 at photoperiods of 12 h or greater. In terms of daily quantum dose, the lowest dose at which greater than 50% gametogenesis occurred was 0.2 E·m^?2·d^?1. However, this critical quantum dose was higher (0.4 E·m^?2·d^?1) when instantaneous irradiances were less than 5 μE·m^?2·s^?1. The saturation quantum dose was also affected by the rate at which the quantum dose was received and varied from 0.4 to 0.8 E·m^?2·d^?1. Gametophytes in all three photoperiods reached 100% fertility at quantum irradiances above 5 μE·m^?2·s^?1. Photoperiod effects were small and could be accounted for by quantum dosage effects.     

TRACE METALS IN GIANT KELP,MACROCYSTIS

Concentrations of 38 elements present in Macrocystis tissues are listed. Mean concentrations of Cu, Zn, Fe, and Mn were similar for samples of juvenile and adult sporophytes from field sites. Cu in Macrocystis tissues rose above background levels when concentrations of 0.05 to 0.1 μm were added to the culturing medium. Above-normal accumulation for Zn occurred at 0.1 μm and for Fe at 10 μm . Tissue Mn appeared independent of the ambient Mn concentration over the ranges studied. Computations indicated that upwelling is important for N and P renewal to Macrocystis beds but may be less significant for import of trace metals.     

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.     

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.     

STUDIES ON THE GIANT KELP,MACROCYSTIS. II. REPRODUCTION

Neushul , M. (U. Washington, Seattle.) Studies on the giant kelp, Macrocystis. II. Reproduction. Amer. Jour. Bot. 50(4): 354–359. Illus. 1963.—The reproduction of Macrocystis pyrifera was studied in the sea and in the laboratory. The estimated minimum time needed for the completion of the sexual life history of Macrocystis pyrifera in La Jolla, California, is from 12 to 14 months. Young sporophytes begin fruiting when they have from 4 to 8 stipes and a somatic frond weight of from 8 to 10 kg. The behavior of abscised sporophylls in laboratory tanks, as well as their morphology and coloration, strongly suggests an interrelationship between the translocation of photosynthetic products and fructification.     

STUDIES ON THE GIANT KELP,MACROCYSTIS. I. GROWTH OF YOUNG PLANTS

Neushul , M., and F. T. Haxo . (Scripps Inst. Oceanography, La Jolla, Calif.) Studies on the giant kelp, Macrocystis. I. Growth of young plants. Amer. Jour. Bot. 50(4): 349–353. Illus. 1963.—The growth of entire young plants of Macrocystis pyrifera was studied in the sea and in the laboratory. Measurements were made of plants in natural stands and of plants transplanted to submerged rafts. Macrocystis was raised in laboratory tanks under controlled conditions of light and temperature. Maximum growth rates in the laboratory paralleled those observed in the sea, plants doubling in area every 16–20 days and in length every 20–30 days under optimum conditions. The transplantation experiments and diving observations demonstrated the important influence of biological activity on the establishment and growth of young plants in the sea.     

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.     

GIANT KELP (MACROCYSTIS PYRIFERA, PHAEOPHYCEAE) RECRUITMENT NEAR ITS SOUTHERN LIMIT IN BAJA CALIFORNIA AFTER MASS DISAPPEARANCE DURING ENSO 1997–1998

During the ENSO event of 1997–1998, density and population structure were evaluated in a Macrocystis pyrifera forest located in Bahía Tortugas, Baja California, Mexico, near the southern limit of the species' distribution in the Northern Hemisphere. Observations in Bahía Tortugas were made quarterly from January 1997 to September 1998 using SCUBA diving surveys. No macroscopic plants were found in the Bahía Tortugas area from October 1997 to April 1998, a local absence of at least 7 months. Aerial surveys further suggest regional disappearance along most of the Baja California coast during the event. Unexpectedly, plants were found in Bahía Tortugas again in July 1998, in spite of the widespread disappearance of the species less than a year earlier. Long-distance spore dispersal was an unlikely cause of the recruitment because: 1) the nearest spore source was more than 100 km away; 2) recruitment appeared to be simultaneous at many sites and occurred rapidly after the cessation of the ENSO event; and 3) the recruits occurred in the same areas as before disappearance. We suggest that a microscopic stage that was not visible during dive surveys survived the stressful conditions of ENSO and caused the recruitment event, supporting the hypothesis that a bank of microscopic forms can survive conditions stressful to macroscopic algae.     

POPULATION SIZE,DISTRIBUTION AND DISPERSAL OF KELP GULLS IN THE SOUTHWESTERN CAPE,SOUTH AFRICA

Steele, W.K. & Hockey, P.A.R. 1990. Population size, distribution and dispersal of Kelp Gulls in the southwestern Cape, South Africa. Ostrich 61:97-106.

There are indications that the Kelp Gull Larus dominicanus population of the southwestern Cape Province, South Africa, is increasing. It is proposed that decreased post-fledging mortality due to supplementary food from man's activities, at sites such as fishing harbours and refuse dumps, is likely to be the main cause of this population increase. Kelp Gulls aggregate, and juveniles (first-year birds) are disproportionately abundant, at sites where supplementary food is available. Juvenile dispersal from the two largest breeding colonies, 38 km apart, is in opposite directions.     

THE ROLE OF DEME SIZE,REPRODUCTIVE PATTERNS,AND DISPERSAL IN THE DYNAMICS OF t-LETHAL HAPLOTYPES

The t-lethal haplotypes (t) found in house mouse (Mus musculus) populations are recessive lethals favored by gametic selection whereby male heterozygotes exhibit a non-Mendelian transmission ratio of about 95% t. The expected equilibrium frequency is 0.385; however, empirical values are lower, averaging close to 0.13. We examined the hypothesis that interdemic selection is the cause of the low empirical values by using a deme-structured simulation model that included overlapping generations, a realistic breeding system, differential deme productivity, and a large total population. We found that under some conditions interdemic selection could lower t frequency below 0.13 in the face of immigration rates up to 5%. Low frequencies were correlated with effective deme size (n_e), regardless of whether n_e was changed through changing deme size (n) or through changing the proportion of breeding adults. Earlier workers showed how the first two phases of interdemic selection (random genetic differentiation and mass selection) interacted to reduce the haplotype frequency, but here we show the importance of the third phase (differential productivity of demes) once demes are linked by dispersal. The effect of this phase is not due to the (negative) covariation between deme productivity and haplotype frequency, but occurs when differential deme productivity generates a difference in t frequency between the population of juveniles recruited into their natal deme and the population of juvenile dispersers. This difference was maximized when the average productivity of demes was low, either because few adult females bred at any one time and/or because fecundity was low. Contrary to an earlier prediction, male-biased dispersal also reduced haplotype frequency, and this probably stems from the relative excess of wild-type genotypes among dispersers compared to the deme residents. Another unexpected finding was that the randomly generated excess of heterozygotes (F_IS < 0) found in small demes favored t haplotypes; however, the effect was only seen when the more powerful influence of the third phase of interdemic selection was removed. Simulations of neutral polymorphisms showed that a deme structure giving F_ST ≤ 0.6 is inconsistent with a haplotype frequency below 0.13. Based on current empirical estimates of F_ST (about 0.2), we concluded that immigration rates in the field are too high for interdemic selection alone to cause the observed deficit of lethal haplotypes. One factor that could combine with population structure effects is the observation that the transmission ratio is lowered to around 0.6 in litters produced from postpartum estrus (PPE). Incorporating this factor, we showed that interdemic selection could be effective in lowering the frequency of t below 0.13 when F_ST was above 0.43 even when migration rates were up to 10%. These results suggest that if empirical haplotype and F_ST estimates are accurate, then additional factors such as a lowered fitness of heterozygotes may be involved.     

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.     

PHOTOSYNTHESIS IN PROTOPLASTS OF MACROCYSTIS PYRIFERA (PHAEOPHYTA)1

Photosynthetic rates measured in protoplasts isolated from the broivn alga Macrocystis pyrifera (L.) Ag. were compared to those for intact tissue. Both ¹⁴C incorporation and O₂ evolution gave similar rates of light-saturated protoplast photosynthesis (approximately 0.4 mmol-g chl a^?1· min^?1). Light saturated photosynthetic rates (P_max) and light harvesting efficiencies (α) of protoplasts were approximately 40% those of intact tissue. In contrast, protoplasts had a greater substrate affinity for photosynthetic HCO₃ uptake (lower K_0.5) than intact tissue (0.87 and 4.1 mMolar, respectively), presumably because of a reduction in the thickness of the unstirred boundary layer in the absence of the cell wall. Overall, the data suggest that protoplasts isolated from Macrocystis pyrifera are of valur in the study of photosynthesis. However, experiments with intact tissue are necessary as controls to aid interpretation of protoplast data.     

GENETIC CONSEQUENCES OF PASSIVE DISPERSAL IN POND-DWELLING COPEPODS

Pond-dwelling copepods have colonized habitats throughout North America after glaciers have receded. Most species are passively transported via resting eggs into new habitats. Colonists originating in a glacial refugium could lose some of the ancestral genetic diversity when they establish new populations and the attenuation may be substantial in populations far removed from the refugium due to multiple founder events. Genetic variation was measured in Heterocope septentrionalis from 27 populations at arctic sites near potential refugia and those more recently deglaciated to determine the effects of postglacial dispersal on patterns of genetic relatedness and diversity. Some populations were more distant, genetically, from others within the same site than those from other more distant sites. Eleven polymorphic enzyme loci were significantly more variable (F [1,294 df] = 5.94, P < 0.025) among individuals from populations near the Alaskan refuge than those at the eastern limit of their distribution. Because populations are typically extremely large and stable this loss of genetic diversity is attributed primarily to repeated founder events during colonization. This result suggests profound genetic changes may occur on a continental scale in passively dispersed copepods due to founder events alone. Their potential for divergence and speciation is greater than currently recognized.     

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.     

CAUSES AND CONSEQUENCES OF WITHIN-TREE PHENOLOGICAL PATTERNS IN THE FLORIDA STRANGLING FIG,FICUS AUREA (MORACEAE)

The obligate pollinators of figs (Ficus, Moraceae), species-specific agaonid wasps, benefit figs only by transporting pollen between trees; larvae are seed predators. But given the high risk of mortality in flight between trees, adult wasps should prefer to pollinate and oviposit within inflorescences (syconia) at the same tree at which they developed. Flowering within individuals is tightly synchronous in most species, while different trees flower out of phase with each other, suggesting that fig phenology has evolved to assure outcrossing. However, some fig species show distinct within-tree flowering asynchrony. It has been suggested that such asynchrony is an adaptation by which figs in seasonal environments can reduce pollinator mortality, by permitting wasps to persist on individual trees at times when flight would be impossible. We have evaluated and rejected the validity of this Seasonality Hypothesis for the Florida strangling fig, Ficus aurea, near its northern range limit. Crops of individual trees were most, not least, synchronous during the coldest, driest months of 2 years. Maximum asynchrony occurred in seasons that were probably most favorable for wasp transit between trees. However, temporal overlap of the phenological stages that permit wasps to remain on their natal trees was always very rare, implying that consecutive cohorts of developing syconia may be spaced in time to limit this occurrence. We suggest alternative costs and benefits for these phenological traits, as well as the proximate mechanisms that might produce them.     

LIFE ON THE EDGE: STRESS SURVIVAL ADAPTATIONS IN SOUTHERN LIMIT MACROCYSTIS PYRIFERA POPULATIONS

In spite of subtropical temperatures often occurring at the Macrocystis pyrifera distributional southern limit in the Northern Hemisphere in Baja California, Mexico, the population has been successful at surviving and recovering from disturbance. To explore the physiological characteristics of southern limit M. pyrifera , microscopic and macroscopic phases of southern limit plants were tested for stress tolerance and compared to northern plant responses in three experiments. In spore development experiments, under constant temperature and nitrate stress treatments, spores from the southern-limit population survived higher temperatures, had greater survivorship, and were able to complete their life cycle at higher temperatures and lower nitrate concentrations than northern population spores. During stress-recovery treatments, where all microscopic stages were tested for recovery after stress shock exposure (temperature, nitrate and light stress), 96% of all haploid treatments showed some regrowth during recovery; haploid survival was not significantly affected by spore population origin, light, or nitrate. However, only 26% of diploid treatments showed regrowth and diploid survival was significantly affected by spore population origin; only the southern limit spores recovered after stress treatment exposure. In juvenile transplant experiments, under all conditions, juvenile plants originating from the southern-limit had higher tissue nitrogen content, greater frond addition rates, and higher photosynthetic efficiency than northern plants, particularly at higher temperatures. The physiological differences observed for the southern limit plants versus those from the north contributes to our understanding of how the southern limit plants can survive in an environment that might otherwise be considered extreme for the species.     

FIELD STUDIES ON THE GIANT KELP NEREOCYSTIS1,2

An intertidal population of Nereocystis luetkeana was investigated in the field, and techniques for measuring sporophyte growth were developed. Primary regions of growth in plants prior to reproductive maturity were located in the bases of the blades, upper stipe, and holdfast. After reproductive areas on blades appeared, growth was confined to 2 major regions: the blade bases and the junction between lower stipe and holdfast where new holdfast haptera were produced. Complete deblading halted growth at whatever stage the plants had attained at the time of the operation but did not kill the plants. Partial deblading resulted in low but measurable growth rates in the blade bases. The data presented suggest that the function of the blades is twofold: they bear reproductive structures in the mature plants and. they may provide, materials used, in rapidly growing regions.