MORPHOLOGICAL VARIATION WITHIN AND BETWEEN NATURAL POPULATIONS OF NON-TIDE POOL FUCUS DISTICHUS (PHAEOPHYTA) IN NEW ENGLAND1,2

The sources of morphological variation in natural populations of Fucus distichus L. ssp. edentatus (De la Pyiaie) Powell and F. distichus L. ssp, evanescens (C. Agardh) Powell from New England were evaluated. Individuals from different populations were morphologically distinct as judged by population means and analysis of variance. A correlation between the plant's form and its habitat was established by field observation. The broadest material grows in calm estuarine habitats, while the narrowest plants occur in exposed, open coastal areas. Major differences in morphology also appear seasonally. In addition, microhabitat factors such as exposure to wave action and elevation explain some morphological variation. Cultured germlings from distinct populations of “evanescens” and “edentatus” type plants were outplanted to an experimental garden in order to ascertain whether the variation was heritable or environmentally induced. The latter experiments indicate that major differences in morphology are genetically determined.     

EFFECTS OF MICROTUBULE INHIBITORS ON PRONUCLEAR MIGRATION AND EMBRYOGENESIS IN FUCUS DISTICHUS(PHAEOPHYTA)1

Pronuclear migration in Fucus distichus spp. edentatus (de la Pyl.) Powell is blocked by incubation of fertilized eggs in colchicine (1 mg/ml) and Nocodazole (2 μg/ ml). Rhizoids form prior to decondensation of the sperm chromatin in eggs in which pronuclear fusion is blocked. This occurs during continuous colchicine incubation as well as in eggs recovering from a short treatment with either drug following fertilization. During recovery of the cells, the sperm and egg chromosomes condense, and the sperm chromosomes migrate toward the egg pronucleus. The delay in migration following removal of colchicine is as much as 24 h and is even slower following removal of Nocodazole. The egg chromosomes form a metaphase plate in treated cells while the sperm chromosomes are still distant in the cytoplasm. This suggests that egg centrioles are important in the mitotic division of the zygote, not sperm centrioles. The effect of colchicine treatment on the mitotic plane and cytokinesis is also discussed.     

VARIABILITY IN TEMPERATURE AND HISTORICAL PATTERNS IN REPRODUCTION IN THE FUCUS DISTICHUS COMPLEX (HETEROKONTOPHYTA; PHAEOPHYCEAE): IMPLICATIONS FOR SPECIATION AND THE COLLECTION OF HERBARIUM SPECIMENS1

Natural and anthropogenic‐induced fluctuations in temperature can have profound effects on an organism's reproduction. The potential of herbarium collections for detecting changes in the timing of reproduction associated with interannual fluctuations in temperature from 1905 to 2003 was assessed in several taxa in the Fucus distichus (L.) emend. Powell complex from northern New England. Data were also used to evaluate whether air and/or sea temperature and North Atlantic Oscillation indices can cause overlap in the reproductive seasonalities of these taxa, a potential mechanism that prevents speciation. For all taxa, there was much variability in reproductive seasonalities among years. Discerning potential correlations with air or sea temperature was difficult because of a suspected bias to collect reproductive specimens at times when most of the population was nonreproductive. Nevertheless, there was a trend for seawater temperature to explain aseasonal reproduction for some taxa. Moreover, the existing archive (1895–2003) shows that the reproductive seasons of these taxa often overlap, and we suggest that interannual variability in temperature may often inhibit speciation. Suggestions also include archive of both reproductive and vegetative materials as well as coordinated sampling in critical biogeographic regions. No one investigator or institution can accomplish this, but phycological societies could organize and sustain such efforts.     

IMMUNOLOCALIZATION OF CENTRIN DURING FERTILIZATION AND THE FIRST CELL CYCLE IN FUCUS DISTICHUS AND PELVETIA COMPRESSA (FUCALES,PHAEOPHYCEAE)1

Antibodies that recognize the centrosome-associated protein centrin were used to characterize centrosomal origin and positioning during fertilization and the first cell cycle in Fucus distichus subsp. evanescens (C. Agardh) Powell and Pelvetia compressa (J. Agardh) De Toni. Centrin was identified in sperm, eggs, and zygotes on protein blots, indicating the protein is present in both gametes. Using immunofluorescence microscopy, centrin was found in discrete foci in sperm. In contrast, eggs lack centrosomes and centrin was not detectable by immunofluorescence, indicating that centrin was probably dispersed in the cytoplasm. Two foci of centrin were present on the nuclear envelope of zygotes, but microtubules remained dispersed over the zygotic nucleus. Centrin foci separated over the nuclear envelope as the first cell cycle progressed. Microtubules became concentrated at the centrin foci to form centrosomes that gave rise to the spindle poles at mitosis.     

EFFECTS OF WOUNDING BY THE HERBIVOROUS SNAILS LITTORINA SITKANA AND L. SCUTULATA (MOLLUSCA) ON GROWTH AND REPRODUCTION OF THE INTERTIDAL ALGA FUCUS DISTICHUS (PHAEOPHYTA)1

Small (3–7 cm long) Fucus distichus ssp. edentatus (de la Pyl.) Pow. Plants were tagged at three sites in which densities of the herbivorous snails Littorina sitkana and L. scutulata ranged from 367–4690 animals · M^-2. From April–August 1986, the growth rate, degree of wounding, and reproductive status of individual thalli were monitored at 2–4 week intervals. Grazer-inflicted damage to the thalli varied within and among sites. Mean growth rates at the site with low densities of littorines were about twice those at the site with intermediate densities and about four times those at the high density site. At the site with high densities of littorines, F. distichus growth rates were negatively correlated with the degree to which the plants were wounded. There appeared to be no correlation of grazer density with F. distichus survivorship. Thalli at the site with few herbivores tended to reproduce earlier and at a larger size than did those at the other two sites. In all three areas, only thalli that had received little damage from herbivores became reproductive. By lowering growth rates and delaying reproduction in F. distichus, grazing by littorine snails can potentially cause variation in reproductive output among individual thalli.     

AN IN VITRO NITRATE REDUCTASE ASSAY FOR MARINE MACROALGAE: OPTIMIZATION AND CHARACTERIZATION OF THE ENZYME FOR FUCUS GARDNERI (PHAEOPHYTA)1

Measurement of the activity of the enzyme nitrate reductase (NR) may provide a useful index of nitrogen metabolism in marine macroalgae. In several species, including Fucus gardneri P. C. Silva, in vitro assays previously failed to detect NR activity, necessitating the use of in situ (or so-called“in vivo”) assays, which are more loosely controlled and lead to dafficulties in assessing enzyme characteristics such as the half-saturation constant (K_m). In this paper, we describe an in vitro NR assay developed for F. gardneri, in which tissue was homogenized using liquid nitrogen prior to the assay. In contrast to previous studies, enzyme activity was always detectable in F. gardneri collected directly from the field at levels up to 30 nmol nitrate converted to nitrite·min^?1·g^?1 wet weight. The effect of a variety of compounds, commonly added to NR extraction buffers, were tested. Additions of protease inhibitors, bovine serum albumin, and ethylenediamine tetraacetic acid had no consistent effects on NR activity, while polyvinyl pyrrolidone, potassium ferricyanide, and flavin adenine dinucleotide significantly decreased activity. The half-saturation constant (K_m) for NADH was 0.18 (± 0.05) mM and for nitrate, K_m=0.99 (±0.41) mM. Significant NR activity was detected without the addition of nitrate, suggesting that internal pools of nitrate averaging approximately 20 μmol NO₃^?·g^?1 wet weight were present in F. gardneri in February. The distribution of NR activity within the plant was highly variable between individuals, but activities were approximately 5-fold lower in the stipe than in midregions. In plants freshly sampled from the field, NR activity increased 7-fold from February to March, then fell to near-February levels by April. These changes in activity may correspond to seasonal changes in growth rate. The assay, optimized for F. gardneri, was used in several different macroalgal species from different taxa: Porphyra sp., Coralina vancouveriensis Yendo, Ulva sp., Enteromorpha intestinalis (Linnaeus) Nees, Macrocystis integrifolia Bory; and Costaria costatum (C. Agardh) Saunders. For all species tested, NR activity was detectable and, except for one species (Porphya sp.), was equal to or greater than activities measured by other workers using in vivo or in vitro assays for plants under similar conditions.     

PHYSIOLOGICAL AND POPULATION ECOLOGY OF INTERTIDAL AND SUBTIDAL ASCOPHYLLUM NODOSUM (PHAEOPHYTA)1

Morphological, demographic and physiological characteristics of Rhode Island intertidal and subtidal populations of Ascophyllum nodosum (L.) Le Jolis were compared in order to examine factors influencing vertical distribution. The two populations had distinctive morphologies: subtidal plants were narrower (more terete) and highly branched compared with intertidal plants. The subtidal population showed signs of necrosis and breakage, which was reflected in significantly shorter mean plant size. High survivorship and low recruitment of both population resulted in relatively constant densities, averaging 91 and 50 plants per m² in the intertidal and subtidal habitats, respectively. Intertidal plants had higher mean annual growth rates (25 cm.yr^?1) than subtidal plants (2 cm.yr^?1). In general, intertidal plants had higher photosynthetic capacity and nutrient (NO₃^?) uptake rates than the subtidal population but maintained lower light-harvesting pigment and tissue nitrogen concentrations. Although Ascophyllum nodosum is capable of survival and growth in subtidal as well as intertidal areas, results of this study suggest that different selective pressures affect persistence in each habitat. The scarcity of plants in the subtidal environment may be due to the lack of a critical balance between algal production, allocation of photosynthate, and the negative effects of grazers or competitors.     

PREMATURE CHROMOSOME CONDENSATION OF THE KARYOGAMY-BLOCKED SPERM PRONUCLEUS IN THE FERTILIZATION OF FUCUS DISTICHUS (FUCALES,PHAEOPHYCEAE)1

Mitosis of egg and sperm pronuclei of Fucus distichus subsp. evanescens (C. Agardh)Powell was examined by fluorescence and electron microscopy when migration of the sperm pronucleus and, as a result, karyogamy were blocked by colchicine treatment after plasmogamy. Chromosome condensation was obsewed in both pronuclei Microspectrophotometric studies after staining the nuclei with mithramycin A clearly showed that DNA synthesis ocurred in the egg pronucleus but not in the sperm pronucleus. This means that chromosomes condensed prematurely in the sperm pronucleus (premature chromosome condensation). In some cases, the egg chromosomes became arranged on a metaphase plate, whereas the sperm chromosomes lay scattered near the egg pronucleus. Immuno fluorescence microscopy using anti-β-tubulin antibody confirmed that a normal spindle was formed at the egg pronucleus. A pair of centrioles existed at the two poles of this spindle. The sperm nuclear membrane disappeared, and microtubules radiated to the sperm chromosomes from one pole of the egg spindle.     

A COMPARISON OF AIR AND WATER AS ENVIRONMENTS FOR PHOTOSYNTHESIS BY THE INTERTIDAL ALGA FUCUS SPIRALIS (PHAEOPHYTA)1

The response of photosynthesis and respiration of the intertidal brown alga Fucus spiralis L. to light and temperature at ambient and elevated concentrations of inorganic carbon was investigated. The light-saturated rate of photosynthesis was greater in air at 15° C and 20° C, but greater in water at 10° C. Light compensation point and I_k was about 50% lower under submerged relative to emerged conditions, whereas the initial slope of photosynthesis versus irradiance was higher, except at 20° C. Under both submerged and emerged conditions light-saturated photosynthesis was limited to a similar degree (78%, and 65%, respectively) by the availability of inorganic carbon at naturally occuring concentrations. In air, slight desiccation at tissue water contents of about 96% to 92% caused a stimulation in the rate of net photosynthesis to 110–148% of fully hydrated fronds. At lower water contents the rate of net photosynthesis declined linearly with decreasing water content and became zero at a water content of about 15%. Dark respiration declined linearly with tissue water content and remained positive to a water content of 8%. Upon reimmersion the fronds showed a complete recovery within 35 min following desiccation to a water content of 20–30%. Thus F. spiralis seems to be very tolerant to desiccation. Since F. spiralis photosynthesizes effectively in air, even at a higher rate than in water as long as it has not lost a large proportion of its water in desiccation, the alternating exposure to air may be beneficial by increasing the daily carbon gain compared to a fully submerged situation.     

SEASONALITY AND THERMAL ACCLIMATION OF REACTIVE OXYGEN METABOLISM IN FUCUS VESICULOSUS (PHAEOPHYCEAE)

The intertidal brown macroalga Fucus vesiculosus L. acclimates its defense against reactive oxygen in response to both (1) growth at different temperatures in laboratory culture and (2) seasonal changes in environmental conditions. Fucus vesiculosus was grown in seawater at 0° C, 20° C, and at 0° C with a 3-h daily emersion at −10° C. Algae grown at low temperature, both with and without freezing, produced less reactive oxygen after severe freezing stress than those grown at 20° C. These differences were correlated with growth temperature-induced changes in activities of superoxide dismutase (SOD), glutathione reductase, and ascorbate peroxidase. The contents of tocopherols increased with increased cultivation temperature, whereas the activity of catalase and the content of glutathione and ascorbate did not change. Growth at 0° C increased the resistance of photosynthesis to freezing and reduced photoinhibition in high light at 5° C; the latter effect was further increased in algae subject to daily freezing. These data suggest that elevated activity of reactive oxygen scavenging enzymes, especially SOD, increases the resistance to photoinhibition, at least at low temperature, as well as being important for freezing tolerance. Seasonal changes in reactive oxygen metabolism showed a similar pattern to those elicited by temperature in laboratory culture. Summer samples had lower activities of most reactive oxygen scavenging enzymes than algae collected in autumn and winter when water temperatures were lower. In contrast to the laboratory experiments, ascorbate content did change and was lower during the winter than summer, whereas the content of glutathione was not influenced by season. Overall, the data not only indicate that temperature plays an important role in the regulation of stress tolerance and reactive oxygen metabolism but also suggest that other factors are also involved.     

A MODEL FOR GAMETE RELEASE IN ASCOPHYLLUM NODOSUM (PHAEOPHYTA)1

The question of what controls gamete release in Ascophyllum nodosum (L.) Le Jolis was studied at six sites along the central coast of Maine. Percent release was assessed weekly along randomly marked transect lines in the mid-intertidal zone. Six independent variables–water temperature at high tide, air temperature at low tide, nitrogen (ammonia, nitrite, and nitrate), and salinity–were measured concurrently. Stepwise multiple regression analysis on the percentage of plants having released gametes revealed that water temperature at high tide accounted for most of the among-site variation (R²= 0.77) in the timing of release. The addition of Julian day increased the R² to 0.82; no other variables were significant. Probit analysis, based on water temperature at high tide, generated an environmentally realistic model for predicting gamete release. The model predicts the onset, midpoint, and termination of gamete release at 6, 10, and 15°C, respectively, and the midpoint at a cumulative water temperature of 358°C. This model has value for developmental studies and, potentially, for reseeding A. nodosum populations. Probits may be useful for characterizing phenological events in other fucoids and algal species.     

INHIBITION OF CARBON UPTAKE AND STIMULATION OF NITRATE UPTAKE AT LOW SALINITIES IN FUCUS DISTICHUS (PHAEOPHYCEAE)1

Inorganic carbon and nitrate uptake were examined in whole plants of Fucus distichus L. (Powell) incubated in dilutions of synthetic ocean water and media with different concentrations of Na⁺, K⁺ and Cl^?. Reduction in salinity from normal seawater (33 ppt) decreased carbon uptake rate but increased nitrate uptake rate by 50% each. Substitution of K⁺ for Na⁺ at constant ionic strength decreased nitrate uptake. Substitution of K⁺ or mannitol for Na⁺ decreased carbon uptake. Neither the uptake of nitrate or carbon was changed by substituting SO₄^2- for Cl^?. Ionophores, valinomycin and monensin, inhibited both nitrate uptake and carbon fixation from 20 to 70% of control rates. The stimulation of nitrate uptake at low salinity may be beneficial to plants in estuarine tidal environments in which nitrate is supplied by the fresh water source.     

PRODUCTION OF HYALINE HAIRS BY INTERTIDAL SPECIES OF FUCUS (FUCALES) AND THEIR ROLE IN PHOSPHATE UPTAKE1

A field study to determine the precise times of year at which three intertidal species of Fucus start to produce hyaline hairs and cease producing such hairs was conducted on the Isle of Man, U.K. Hairs were first observed during February, and within 6 days of their initial appearance, all tagged plants of all species at all tidal heights on the shore possessed hairs. Hair production continued until the beginning of October, at which time Fucus plants growing at the lowest stations (+ 3.0 m) had glabrous apical growth. Hair production continued later into the year for plants growing higher on the shore, and it was not until mid-November that glabrous apical growth was observed in all plants. Phosphate uptake rates of pilose (hairy) and glabrous (hairless) apical sections were measured in November 1988 for F. spiralis L. and in January 1989 for F. spiralis and F. serratus L., at phosphate concentrations ranging from 0.8 μM (ambient seawater) to 9.0 μM. In ambient seawater, pilose plants of F. spiralis removed phosphate 2–3 times faster than glabrous plants, whereas the uptake rates of pilose plants of F. serratus were about 50% greater than those of glabrous plants. The differences between uptake rates of pilose and glabrous plants of both species were smaller or nonsignificant at higher phosphate concentrations. The field and laboratory data are consistent with the hypothesis that hairs are formed in Fucus as a response to increased nutrient demand and that hairs facilitate the uptake of nutrients from seawater at concentrations typical of natural situations.     

A NEW, NONRADIOACTIVE WHOLE-MOUNT IN SITU HYBRIDIZATION PROTOCOL FOR FUCUS (PHAEOPHYTA) EMBRYOS

We present a nonradioactive protocol to localize mRNA in Fucus distichus ssp . edentatus ( de la Pyl.) Powell (Phaeophyta) embryos using whole-mount in situ hybridization. Hybridization of a digoxygenin (DIG)-labeled oligo-dT probe to poly A⁺ RNA, and DIG-labeled riboprobes to specific mRNAs, was detected with an antibody to DIG (anti-DIG) that was coupled to alkaline phosphatase. Our protocol, a modification of one developed for higher plants, includes a treatment of fixed embryos with specific cell-wall-degrading enzymes to permeabilize the cells to anti-DIG, progressive osmotic changes to prevent plasmolysis, and a RNase A treatment to reduce nonspecific background. Using this protocol, we show the intracellular distribution of poly A⁺ RNA and mRNAs for actin and the fucoxanthin chlorophyll protein A (Fcp A). This method to detect mRNA localization is rapid because it does not require embedding, sectioning, or the use of radioactivity. With the availability of a variety of enzymes to degrade plant cell walls, this protocol should be applicable to many algal cells .     

REACTIVE OXYGEN PRODUCTION AND DAMAGE IN INTERTIDAL FUCUS SPP. (PHAEOPHYCEAE)

The research described in this paper was designed to test the hypothesis that the differential stress tolerance associated with the vertical zonation of intertidal seaweeds is attributable to reactive oxygen metabolism. To do so, we measured the production of, and damage caused by, reactive oxygen in three species of intertidal brown seaweeds— Fucus spiralis L., F. evanescens C. Ag., and F. distichus L.— that differ in their ability to withstand freezing, desiccation, and high light stress. Fucus spiralis is the most stress-tolerant species and F. distichus the least. Reactive oxygen production was determined by measuring the production of H₂O₂ and the oxidation of dichlorohydrofluorescein diacetate to dichlorofluorescein. Damage caused by freezing, desiccation, and high-light stress was assessed by measuring variable fluorescence (F_v/F_m) and lipid peroxidation. Production of reactive oxygen increased following freezing, desiccation, or high-light stress. In general, the data were consistent with the hypothesis that reactive oxygen metabolism is involved in stress tolerance. The production of reactive oxygen was relatively low in unstressed seaweeds, and there was little difference between species. Fucus distichus showed the greatest increase in reactive oxygen production after desiccation and freezing stress. Fucus evanescens produced more reactive oxygen production after desiccation than F. spiralis. Although F. evanescens and F. spiralis produced similar amounts of reactive oxygen after freezing, this treatment resulted in an increase in lipid peroxidation only in F. evanescens (and F. distichus ).     

DETERMINING THE AFFINITIES OF SALT MARSH FUCOIDS USING MICROSATELLITE MARKERS: EVIDENCE OF HYBRIDIZATION AND INTROGRESSION BETWEEN TWO SPECIES OF FUCUS (PHAEOPHYTA) IN A MAINE ESTUARY1

The high degree of morphological plasticity displayed by species of the brown algal genus Fucus L. is well documented. Such variation is especially pronounced for those estuarine taxa lacking holdfasts (termed ecads) that often bear little resemblance to the attached species from which they are derived. To better understand the systematics of salt marsh fucoids, we developed a suite of four microsatellite‐containing loci capable of distinguishing between F. vesiculosus L. and F. spiralis L. The genetic markers were used to determine the relationships of the fucoid ecads F. vesiculosus ecad volubilis (Hudson) Turner and a muscoides‐like Fucus in the Brave Boat Harbor (ME, USA) estuary. Ecad populations had 2‐ to 3‐fold higher levels of heterozygosity than attached populations of F. vesiculosus and F. spiralis. Further, ecads were “intermediate” between F. vesiculosus and F. spiralis in their allele frequencies and genotype composition. Our data indicate that populations of muscoides‐like Fucus in Brave Boat Harbor mainly consist of F₁ hybrids between F. vesiculosus and F. spiralis, whereas F. vesiculosus ecad volubilis may arise through backcrosses between F. vesiculosus and other fertile hybrids. Finally, our data support the hypothesis that introgression has occurred between attached populations of F. vesiculosus and F. spiralis.     

FUNCTIONAL-MORPHOLOGICAL RELATIONSHIPS IN SARGASSUM POLYCERATIUM (PHAEOPHYTA): PHENOTYPIC AND ONTOGENETIC VARIABILITY IN APPARENT PHOTOSYNTHESIS AND DARK RESPIRATION1

Phenotypic and ontogenetic changes in apparent photosynthesis and dark respiration for Sargassum polyceratium Mont. were related to the differential allocation of resources into blades, stems, vesicles, and fertile branches as well as anatomical parameters of surface area, volume, and blade density from a population in the Content Keys, Florida. Three divergent phenotypes were evaluated during four phases of ontogeny: maximum growth (October), peak reproduction (December), senescence (April), and stasis (July). Photosynthetic and respiratory rates changed throughout the year, at different stages of morphogenesis, and among different phenotypes. Photosynthesis was highest during active growth, decreased during reproduction and senescence, and was lowest during stasis. In contrast, respiration was higher during peak reproduction and stasis. Temporal, phenotypic, and ontogenetic changes in apparent photosynthesis were best explained by interactions among anatomical features, growth stages, and source-sink relationships of metabolic reserves and pigments. Surface area: volume (SA:V) ratios played a secondary role. Photosynthetic performance of mature axes decreased with the allocation of resources away from blades to air bladders and fertile branches. Apparent productivity and development of fertile branches on phenotypes were not correlated. High respiratory rates, SA:V ratios, blade areas, and low blade densities occurred on developing axes and were consistent with adaptations to low light fields. Sargassum polyceratium, because of its morphological differentiation, is a relatively complex, physiologically variable alga.     

THE EFFECT OF INTERTIDAL STRESS ON PADINA SANCTAE-CRUCIS (PHAEOPHYTA)1

The hypothesis is that plants growing in the intertidal region display developmental and morphological differences from subtidal plants of the same species. Intertidal and subtidal populations of Padina sanctae-crucis Börg. Were compared, emphasizing morphological and developmental features. Plant size in relation to number of reproductive bands, vegetative cell sizes and the rate of spore development were analyzed to evaluate the hypothesis. The results indicate that environmental stress tends to dwarf intertidal plants through reduction in cell size. Spores from both intertidal and subtidal populations were cultured in the laboratory under standard conditions. No significant differences were found between the growth rates. The results may be interpreted to mean that cell enlargement is inhibited by intertidal stress.     

EVIDENCE FOR METHYL JASMONATE-INDUCED PHLOROTANNIN PRODUCTION IN FUCUS VESICULOSUS (PHAEOPHYCEAE)

The plant growth regulators jasmonic acid and methyl jasmonate (MeJA) have recently been identified in a variety of marine algae; however, their role in these organisms is currently unknown. Here we report that exposure to MeJA, during periods of tidal emergence causes the induction of polyphenolic chemical defenses (the phlorotannins) in two populations of the common rockweed Fucus vesiculosus (Linnaeus). Phlorotannin concentrations were up to 1.6 times higher in the growing apices of F. vesiculosus from both Avery Point (Connecticut, USA) and Roosevelt Inlet (Delaware, USA) within 10–14 days after a single brief exposure to airborne MeJA at concentrations ranging from 5.42 to 542 nM. The timing and magnitude of this induced increase in phlorotannin concentration are similar to that caused by real and simulated herbivory, raising the question of whether jasmonates, or their oxylipin relatives, are natural elements of antiherbivore responses in Fucus , as they are in vascular plants.     

CRYOANALYTICAL STUDIES OF FREEZING DAMAGE AND RECOVERY IN FUCUS VESICULOSUS (PHAEOPHYCEAE)

Receptacles of Fucus vesiculosus L. were examined using cryoscanning electron microscopy of rapidly frozen, hydrated tissue to avoid extraction and collapse of the extracellular matrix (ECM) and provide new structural information. The pore of the conceptacle was occluded by the tightly appressed tips of the paraphyses. These were embedded in copious ECM within the conceptacle. The ECM inside the conceptacle and in the medulla differed markedly in texture and with respect to the distribution of S and Cl, as revealed by energy-dispersive X-ray microanalysis. High levels of S were restricted to the conceptacle's ECM, whereas the ECM in the medulla was always S poor and Cl rich. Receptacles frozen naturally at low tide during the winter of 1998 showed extensive damage in the medulla, and this likely accounts for the cessation of reproduction during the winter on the Maine shore. The damaged receptacles recovered and resumed growth during the spring, and they released gametes by May. Abscission of the overwintered receptacles was observed in June to July, coincident with the formation of new receptacles at the tips of newly grown portions of the fronds.