PREDICTING DESICCATION STRESS IN MICROSCOPIC ORGANISMS THE USE OF AGAROSE BEADS TO DETERMINE EVAPORATION WITHIN AND BETWEEN INTERTIDAL MICROHABITATS1

We describe an easy and inexpensive way to determine whether intertidal microhabitats remain wet during tidal emersion. This new technique uses agarose beads (120 üm diameter when fully hydrated) that shrink in a graded fashion as they dry. The agarose beads allow variability in surface wetness to be gauged over distances of less than 1 mm. Describe this parameter of microclimate is important in order to predict the likelihood and spatial pattern of survival of settled larvae, reproductive propagules, and other microscopic stages in the life histories of organisms growing in intertidal and other water-stressed environments. For the brown seaweed Pelvetia fastigiata (J. Ag.) DeToni (Fucales, Phaeophyta), the use of agarose beads demonstrated that survival of zygotes during tidal emersion was highes at those sites that remain damp. Temperature alone was found to be an unreliable measure of wetness within a single microhabitat (e.g. red algal turf).     

ION FLUXES AND MODIFICATION OF THE EXTRACELLULAR MATRIX DURING GAMETE RELEASE IN FUCOID ALGAE

A rapid phase transition in the extracellular matrix (ECM) of the reproductive tissues (= receptacles) has been proposed to cause gamete release in fucoid algae. We tested this model with cryoanalytical energy dispersive x-ray microanalysis (EDX) scanning electron microscopy (SEM) of rapidly frozen hydrated receptacles of Silvetia compressa Serrão, Cho, Boo et Brawley that were planed to smooth faces using a cryoultramicrotome. Every receptacle typically contained a region(s) of intracellular accumulation of K and Cl and a region(s) of efflux of K and Cl to the ECMs, regardless of treatment (e.g. calm vs. shaken conditions in the light, anthracene-9-carboxylic acid). Although longitudinal variations in [K] and [Cl] between ECM and cells were common, a tissue within any transverse plane of the receptacle was in the same state of efflux or accumulation of K and Cl. The strongest and most extensive effluxes occurred during the time of irreversible commitment to gamete release (i.e. 2–4 min darkness after light potentiation). A receptacle may oscillate rapidly between global states of accumulation and efflux at this time. The polysaccharide collar between the oogonium and stalk cell expanded during Cl and K efflux into the ECM of the conceptacle and condensed when [K] and [Cl] were low in the conceptacle ECM. During periods of strong efflux associated with gamete release, the collar and other components of the exochiton ruptured. Detached gametangia were guided to the pore by the paraphyses. The ECM of the conceptacle is sulfur rich (226 mM) compared with the ECM of the medulla (8 mM). This analysis supports the importance of osmotic modification of the ECM during gamete release and demonstrates that the receptacle is a dynamic signaling organ.     

PHOTOSYNTHESIS,DARK RESPIRATION AND DESICCATION RESISTANCE OF THE INTERTIDAL SEAWEEDS HESPEROPHYCUS HARVEYANUS AND PELVETIA FASTIGIATA F. GRACILIS12

Rates of net photosynthesis and dark respiration were determined under submersed and emerged conditions for Hesperophycus harveyanus S. & G. and Pelvetia fastigiata f. gracilis (Decne.) S. & G. Both species exhibited submersed photosynthesis-light relationships and dark respiration rates similar to those established for other closely related intertidal, fucoids. Maximal net photosynthesis of H. harveyanus (0.21 mmol O₂ g dry wt.^-1· h^-1; 0.18 mmol CO₂ g dry wt.^-1· h^-1) was similar to that of P. fastigiata f. gracilis (0.17 mmol. O₂ g dry wt.^-1· h^-1; 0.14 mmol CO₂ g dry wt. ^-1· h^-1). Light saturation occurred between 150 and 250 μE · m^-2· s^-1 for H. harveyanus and between 75 and 150 μE · m^-2· s^-1 for P. fastigiata f. gracilis; photon flux densities required for compensation were 6.4 and 9.2 μE · m^-2· s^-1, respectively. Photoinhibition was not observed for either species. The light-saturated, submersed net photosynthetic performances of both species varied significantly with temperature. Greatest photosynthetic rates were obtained at 23° C for H. harveyanus and at 18° C for P. fastigiata f. gracilis. Under emersed conditions, the maximal net photosynthetic rate and the photon flux densities required for saturation were greater for H. harveyanus (0.08 mmol CO₂ g dry wt.^-1· h^-1; 260 to 700 μE · m^-2· s^-1) than for P. fastigiata f. gracilis (0.02 mmol CO₂g dry wt.^-1· h^-1; 72 to 125 μE · m^-2· s^-1). However, for both species, emersed photosynthetic rates were much lower (14–44%) than those obtained under submersed conditions. Desiccation negatively influenced emersed photosynthesis, of both species, but H. harveyanus thalli contained more water when fully hydrated and lost water more slowly during dehydration, thus suggesting greater photosynthetic potential during field conditions of emersion.     

COMPONENTS OF PHOTOSYNTHESIS IN THE INTERTIDAL SACCATE ALGA HALOSACCION AMERICANUM (RHODOPHYTA,PALMARIALES)1

Rates of photosynthesis for the intertidal saccate alga Halosaccion americanum Lee were determined under submersed and emersed conditions. By fitting the data to a hyperbolic tangent function, P _max was 4.08 mmol CO₂. m^?2. h^?1 and I_k was 116.4 μE. m^?2. s^?1. under submersed conditions. Under emersed conditions, P _max was 1.89 mmol CO₂. m^?2. h^?1 and I_k was 22.9 μE. m^?2. s^?1. Dark fixation represented 3.7% of P_max in submersed thalli, whereas it equalled 33.3% of P_max in emersed thalli. Photosynthetic uptake from the thallus cavity represented a significant source of carbon, achieving 68.8% of that from the atmosphere and 29.4% of that from seawater. Retained seawater also greatly reduced drying under emersed conditions. Experimental thalli lost 70.4% of their water after 120 min under desiccating conditions, whereas control thalli lost only 6.3%. Emersed photosynthetic rates were enhanced by desiccation, At times, rates for desiccated thalli were two times those of fully-hydrated ones. Only after water loss exceeded 47% did photosynthetic rates fall below fully-hydrated rates. Utilizing data from this study a model was constructed to determine total photosynthetic production of H. americanum over a single daylight period. These caluclations demonstrate that photosynthetic contributions from emersed photosynthesis and retained seawater are significant. Because production from all sources is almost equal, total photosynthesis over a single day does not change greatly regardless of the time spent in air or in water.     

MACROALGAL RECRUITMENT IN A HIGH MARSH CREEK OF NORTH INLET ESTUARY,SOUTH CAROLINA1

The temporal and spatial variability in recruitment patterns of macroalgae were stuided by evaluating the appearace of propagules on marble tiles set over a depth gradient in a high marsh tidal creek in North Inlet Estuary, South Carolina. Ulvoids and Porphyra occurred over a wide range of depths in the intertidal zone unlike Ectocarpus and members of the Florideophycease, which grew over a more restricted zone. Ulvoid propagules were the most abundant, attaining maximum densities at 0 and -15 cm mean low water (MLW) at all seasons except winter. In summer, coincident with high tubbidity, ulvoid densities decreased at -15 cm MLW but not in other seasons. Ulvoid densities at 0 MLW were depressed in January, February was preceded with the lowest recoreded temperatures and salinities during the study period, while the decrease in July was preceded by the highest recorded temperature. The number of taxa was highest in April and September, representing winter-spring and summer-fall transition periods, respecitively. The biomass of adult forms was higher on the tiles than on naturally occurring substrala, perhaps due to lack of stable suitable substrata in the field. Inhibition of subsequent recruitment by initial recreuitment was evident only on tiles submerged in October. The initial recruitment pattern of Porphyra rosengurtii coll et Cox and Ectocarpus siliculosus (Dillwyn) Lyngbye seem to determine the biomass distribution of adult forms, while that of ulvoids may be altered later by other factors. The considerable decrease in the number of adult forms compared to the initial denisties of propagules indicales high juvenile mortalities.     

THE IMPACT OF FROND CROWDING ON FROND BLEACHING IN THE CLONAL INTERTIDAL ALGA MAZZAELLA CORNUCOPIAE (RHODOPHYTA, GIGARTINACEAE) FROM BRITISH COLUMBIA, CANADA

The importance that frond crowding represents for the survival of fronds of the clonal intertidal alga Mazzaella cornucopiae (Postels et Ruprecht) Hommersand (Rhodophyta, Gigartinaceae) was investigated in Barkley Sound, British Columbia, Canada. Frond density is high for this species, up to 20 fronds·cm^?2 in the most crowded stands. Frond crowding imposes a cost in the form of reduced net photosynthetic rates when fronds are fully hydrated as a result of reduced irradiance compared with experimental (not found naturally) low-density stands. However, the interaction between desiccation and irradiance alters this relationship between net photosynthetic rates and frond density. During a typical daytime low tide in spring, irradiance is 10–30 μmol·m^?2·s^?1 below the canopy of fronds, and frond desiccation (relative to total water content) can reach 43% at the end of the low tide. In contrast to natural stands, fronds from experimentally thinned stands are subjected to irradiances up to 2000 μmol·m^?2·s^?1 because of the spatial separation among fronds and can desiccate up to 81% at the end of the same low tide. Laboratory experiments showed that negative net photosynthetic rates occur between 40% and 80% desiccation at an irradiance of 515 μmol·m^?2·s^?1, and the literature suggests that strong bleaching could occur as a result. At 20 μmol·m^?2·s^?1 of irradiance and desiccation levels up to 40%, simulating understory conditions of natural stands, net photosynthetic rates are never negative. Experimental thinning of stands of M. cornucopiae done during spring effectively resulted in a stronger extent of frond bleaching compared with natural stands. Therefore, the cost of reduced net photosynthetic rates at high frond densities when fronds are fully hydrated is counterbalanced by the protective effects of frond crowding against extensive bleaching, essential for survival at the intertidal zone. Future research will have to demonstrate the possible relationship between the frequency and duration of negative net photosynthetic rates and the extent of frond bleaching.     

POPULATION STRUCTURE AND PHYSIOLOGICAL DIFFERENTIATION OF HAPLOTYPES OF CALOGLOSSA LEPRIEURII (RHODOPHYTA) IN A MANGROVE INTERTIDAL ZONE

Zonation of macroalgae in the intertidal zone has been well documented. However, studies of zonation of macroalgae have predominantly examined the distribution of different species rather than the distribution of variants within a species. This study investigated the spatial variation of plastid haplotypes of the mangrove red alga Caloglossa leprieurii (Montagne) J. Agardh at a site in eastern Australia and tests for physiological differences (growth, photosynthesis) between those haplotypes. RUBISCO spacer plastid haplotypes were scored using single-stranded comformational polymorphism, and the population structure at two sites was examined using a nested sampling design comparing between sites, among transects within sites, and among quadrats within transects. Growth rates at various salinities and light intensities and the photosynthesis–irradiance curves of the three main haplotypes were compared. The two sites showed a high degree of genetic differentiation across a short distance, suggesting limited gene flow. The distribution of haplotypes was patchy and did not reflect a zonation pattern along the intertidal gradient. The three haplotypes were physiologically differentiated with haplotype A, with a lower growth rate and a lower photosynthetic efficiency at higher light intensities. There is some evidence of physiological differentiation between life history phases in C. leprieurii with sporophytes having a higher growth rate than females under most conditions. Our results suggest a correlation between our culture results and our population data. Haplotypes (haplotype A) and life history phases (gametophytes) with lower performance (growth and photosynthetic efficiency) under our culture conditions were correlated with a minor representation in the field. This is the first study to integrate population-level data with physiological parameters toward an understanding of the distribution and relative abundance of red algal genetic variants.     

INVOLVEMENT OF MICROCYSTINS AND COLONY SIZE IN THE BENTHIC RECRUITMENT OF THE CYANOBACTERIUM MICROCYSTIS (CYANOPHYCEAE)1

The benthic recruitment of Microcystis was simulated in vitro in order to characterize the colonies of Microcystis recruited and to study the impact of intracellular and extracellular microcystins (MCs), and the influence of colony size on the recruitment process. We observed recruitment dynamics consisting of a lag phase followed by a peak and then a return to low recruitment rates, mainly controlled by passive resuspension throughout the experiment, and by physiological processes during the recruitment peak. Ninety‐seven percent of the Microcystis colonies recruited were <160 μm in maximum length, and their cells contained much greater amounts of MCs (0.26 ± 0.14 pg eq microcystin leucine‐arginine variant [MC‐LR] · cell^?1) than those in benthic colonies (0.021 ± 0.004 pg eq MC‐LR · cell^?1). The MC content of recruited Microcystis varied significantly over time and was not related to changes in the proportion of potentially toxic genotypes, determined using real‐time PCR. On the other hand, the changes in MC content in the potentially toxic Microcystis recruited were closely and negatively correlated with recruitment dynamics; the lowest MC contents corresponded to high recruitment rates, and the highest MC contents corresponded to low recruitment rates. Thus, depending on temperature and light conditions, these variations are thought to result from the selection of various subpopulations from among the smallest and the most toxic of the initial benthic population. Adding purified MC‐LR to experimental treatments led to a decreased recruitment of Microcystis and more specifically of mcyB genotypes.     

PHYSIOLOGICAL SNAPSHOTS REFLECT ECOLOGICAL PERFORMANCE OF THE SEA PALM,POSTELSIA PALMAEFORMIS (PHAEOPHYCAEA) ACROSS INTERTIDAL ELEVATION AND EXPOSURE GRADIENTS1

Postelsia palmaeformis Ruprecht is an intertidal kelp found only on very wave‐exposed rocky shores of the northeast Pacific. In areas dominated by mussels, Postelsia depends on wave‐induced disturbances to complete its life‐history cycle. Postelsia also recruits where mussels are absent, but not at less wave‐exposed shores. Thus, physical conditions related to wave exposure limit its horizontal distribution. It is not clear what limits the vertical distribution of Postelsia. We investigated factors contributing to Postelsia's limited distribution using transplant experiments, demographic monitoring, and field fluorometry to evaluate growth and performance across gradients of tidal elevation and wave exposure. Survivorship and growth were sharply reduced at upper and wave‐protected edges relative to mid‐level, wave‐exposed sporophytes. Reproductive output was reduced at upper and lower levels, and growth but not survivorship was lower at the lower level. Effects were independent of population of origin and were a manifestation of the environment. Maximum electron transport rates (ETR_m), light saturation parameters (E_k), and maximum quantum yields (ΔF/F_m) provided insight into physiological dynamics; all were lowest at the high edge, but increased when desiccation stress was alleviated by a mock sea‐spray treatment. The ETR_m and E_k values of low sporophytes were not as high as the values for mid‐sporophytes, despite higher or equivalent nitrogen content, chl a, and absorptance, suggesting a trade‐off between light‐capturing and carbon‐fixation capacity. Physiological limitations at upper and lower levels and deleterious desiccation effects at wave‐protected sites prevent establishment, thus constraining Postelsia to a mid‐zone, wave‐exposed distribution. Physical conditions related to wave exposure may limit the horizontal distribution of Postelsia because this kelp is also found in areas where mussels are lacking but not on less wave‐exposed shores.     

REPRODUCTION AND MORPHOLOGICAL VARIATION IN SOUTHERN CALIFORNIA POPULATIONS OF THE LOWER INTERTIDAL KELP EGREGIA MENZIESII (LAMINARIALES)1

Intertidal Egregia menziesii (Turner) Aresch. populations were studied at three Southern California sites to determine temporal and spatial patterns of reproduction and morphology. The timing of sporophyll production and sporophyte recruitment was similar at all sites. Sporophyll production was much greater during winter periods of colder seawater temperatures and shorter day lengths. Sporophyte recruitment occurred from spring through midsummer, ～5 months following maximal sporophyll production. Lateral blade morphologies varied in a consistent manner, suggesting a developmental mechanism for form variation in Egregia thalli. Spatulate blades dominated shorter axes and the bases of longer axes, whereas filiform laterals became abundant toward the tips of longer axes. Filiform laterals (9.8 mg O₂·g^?1·h^?1) had higher light‐saturated net photosynthetic rates than spatulate laterals (6.8 mg O₂·g^?1·h^?1), resulting in a 12% increase in the productivity of Egregia per meter of filiform frond.     

ROLE OF SURFACE WOUNDS AND BROWN ALGAL EPIPHYTES IN THE COLONIZATION OF ASCOPHYLLUM NODOSUM (PHAEOPHYCEAE) FRONDS BY VERTEBRATA LANOSA (RHODOPHYTA)1

Ascophyllum nodosum (L.) Le Jol. forms extensive beds in wave‐sheltered, rocky intertidal habitats on the northwestern Atlantic coast. This fucoid seaweed is host to an obligate red algal epiphyte, Vertebrata lanosa (L.) T. A. Chr. [=Polysiphonia lanosa (L.) Tandy], and two facultative brown algal epiphytes, Elachista fucicola (Velley) Aresch. and Pylaiella littoralis (L.) Kjellm. Although V. lanosa can occur throughout most of the length of host fronds, it largely predominates in midfrond segments. The two brown algal epiphytes are restricted to distal segments. Through field experiments conducted in Nova Scotia, Canada, we tested the hypothesis that surface wounds are required for the colonization of distal segments of host fronds by V. lanosa. Distal tissues normally have a smooth surface because of their young age (A. nodosum fronds grow apically). By creating small wounds that mimicked grazing wounds distributed elsewhere on host fronds, we demonstrated that V. lanosa can colonize distal frond segments during the growth and reproductive season (summer and autumn). Approximately half of the artificial wounds were colonized by V. lanosa during this time. The experimental exclusion of both brown algal epiphytes from distal frond segments did not affect colonization by V. lanosa. Thus, we conclude that the absence of surface irregularities on distal segments of host fronds, specifically small wounds, is the main factor explaining the absence of V. lanosa there. We propose that further experimental work clarifying epiphyte distribution in host beds will enhance our ability to understand the functional role of epiphytes in intertidal ecosystems.     

俄罗斯平原针阔混交林群落的灌木层植物种间相关研究

灌木层植物种类丰富,成层明显,结构复杂是俄罗斯平原针阔混交林群落的突出特点,应用Ｘ＾２检验和Ｐｅａｒｓｏｎ相关系数ｒ公式研究了灌木层植物种间联结和相关性,对测定结果的生态学意义进行了分析,依据测定结果,将形成灌木层的植物划分为４个生态联结种组;俄罗斯金雀花＋璎珞柏,欧洲白蜡＋欧洲白榆,疣枝卫矛＋欧洲荚迷＋黄忍冬＋欧榛子＋黑茶镳子＋欧洲赤杨。生态种组的划分为整个群落的结构和动态研究及植被的恢复和重建     

EFFECTS OF CURRENT VELOCITY AND LIGHT ENERGY ON THE STRUCTURE FO PERIPHYTON ASSEMBLAGE IN LABORATORY STREAMS1

Effects of current velocity and light energy on the taxonomica and physiognomic characteristics of periphyton assemblages were investigated in laboratory streams. The initial rate of colonization was related to current velocity, while the effects of light energy accounted for differences in species composition by the end of the experiment. Although the laboratory systems had many species in common during the realy stages of colonization, the experimental treatments generated differences in rates of communitydevelopment. synedra spp. were the early coloniters of the substrate, followed by an understory of Achnanthes spp. After day 16, Stigeoclonium tenue developed in the streams exposed to the higher photon flux density, but was rare in the shaded streams. The applicability of traditional successional theory to develoopmental patterns in lotic periphyton assemblages is discussed.     

KELP PATCH DYNAMICS IN THE FACE OF INTENSE HERBIVORY: STABILITY OF AGARUM CLATHRATUM (PHAEOPHYTA) STANDS AND ASSOCIATED FLORA ON URCHIN BARRENS1

The brown alga Agarum clathratum (Dumortier) is the only large, perennial, fleshy macrophyte commonly found on urchin‐dominated barrens in the northwestern North Atlantic. We examined the spatial and temporal stability of A. clathratum stands and their impact on algal recruitment in the Mingan Islands, northern Gulf of St. Lawrence. The stands were highly stable in space and time, with only small intersite variations. The percent cover of A. clathratum in 144‐m² areas increased by 6.5%–11.4% over a 2‐year period, and most changes in abundance occurred at the edge of the stands. The surface area of small (<13 m²) single stands of A. clathratum increased by approximately 1.8%·month^?1, although marked increases (>95%) occurred during winter, largely because adjacent stands merged into larger single stands. Mature stands of A. clathratum appear to enhance algal recruitment, as juvenile A. clathratum and the understory red alga Ptilota serrata (Kützing) were orders of magnitude more abundant inside than outside the stands. The experimental removal of the A. clathratum canopy (1‐m² portions) had no long‐term effect on the abundance of A. clathratum, which within 14 months had recolonized most of the cleared areas. In contrast to juvenile A. clathratum, the abundance of P. serrata rapidly decreased after canopy removal. Our results demonstrate that A. clathratum stands are a stable component of urchin barrens in spite of the heavy grazing that typically occurs there. Maintenance and expansion of A. clathratum stands and associated flora appear to depend on positive interactions with self‐defended adult A. clathratum.