High-temperature tolerance of photosynthesis in the red alga Chondrus crispus

Chondrus crispus (Stackhouse) is a perennial red seaweed, common in intertidal and shallow sublittoral communities throughout the North Atlantic Ocean. In the intertidal zone, C. crispus may experience rapid temperature changes of 10 to 20C° during a single immerison-emerision cycle, and may be exposed to temperatures that exceed the thermal limits for long-term survival. C. crispus collected year-round at Long Cove Point, Chamberlain, Maine, USA, during 1989 and 1990, underwent phenotypic acclimation to growth temperature in the laboratory. This phenotypic acclimation enhanced its ability to withstand brief exposure to extreme temperature. Plants grown at summer seawater temperature (20°C) were able to maintain constant rates of lightsaturated photosynthesis at 30°C for 9 h. In contrast, light-saturated photosynthetic rates of plants grown at winter seawater temperature (5°C) declined rapidly following exposure to 30°C, reached 20 to 25% of initial values within 10 min, and then remained constant at this level for 9 h. The degree of inhibition of photosynthesis at 30°C was also dependent upon light intensity. Inhibition was greatest in plants exposed to 30°C in darkness or high light (600 mol photons m^-2s^-1) than in plants maintained under moderate light levels (70 to 100 mol photons m^-2s^-1). Photosynthesis of 20°C-acclimated plants was inhibited by exposure to 30°C in darkness or high light, but the degree of inhibition was less than that exhibited by 5°C-grown plants. Not only was light-saturated photosynthesis of 20°C plants less severely inhibited by exposure to 30°C than that of 5°C plants, but the former also recovered faster when they were returned to growth conditions. The mechanistic basis of this acclimation to growth temperature is not clear. Our results indicate that there were no differences between 5 and 20°C-grown plants in the thermal stability of respiration, electron transport associated with Photosystems I or II, Rubisco or energy transfer between the phycobilisomes and Photosystem II. Overall, our results suggest that phenotypic acclimation to seawater temperature allows plants to tolerate higher temperatures, and may play an important role in the success of C. crispus in the intertidal environment.     

Inorganic carbon uptake by photosynthetically active protoplasts of the red macroalga Chondrus crispus

Photosynthetically active protoplasts were isolated from Chondrus crispus Stackh. by treating thalli with -carrageenase produced from batch culture of Pseudomonas carrageenovora. Using the silicone oil centrifugation technique, it was found that the protoplasts: (1) did not generally accumulate inorganic carbon (C_i) above the concentration in their incubation medium; (2) were saturated at C_i concentrations of 3 to 4 mM; (3) had an intracellular pH of 7.50 when incubated at pH 7.5; and (4) their initial carbon fixation rate was reduced by carbonic anhydrase inhibitors. Although the carbon fixation rate of the protoplasts was about 30% that of thallus fragments, presumably due to the relatively harsh protoplast isolation treatment, the behavior of the protoplasts was similar to that of fragments. This similarity indicates that the protoplasts are photosynthetically active and behave as thallus fragments. Further, the data are consistent with the hypothesis that C. crispus acquires C_i for photosynthesis by the diffusion of CO₂ across the plasma membrane.     

Temperature-influenced infection rates in the Chondrus crispus—Petersenia pollagaster pathosystem: a regression analysis

Cross-infection experiments were performed to determine the influence of temperature on infection rate in the Chondrus crispus Stackhouse-Petersenia pollagaster (Petersen) Sparrow pathosystem. C. crispus thalli were collected at Pubnico Harbor, Nova Scotia, Canada in the fall of 1981 to 1984. Infective zoospores were used to inoculate healthy thalli at five different temperatures. The highest infection rate was obtained at 20°C, while significantly lower rates were obtained at temperature extremes. The parasite's life cycle, consisting of infection of healthy thalli, endobiotic development, and release of zoospores, was completed in 48 to 72 h at 15° to 20°C.     

Effects of high-frequency light fluctuations on growth and photoacclimation of the red algaChondrus crispus

High-frequency fluctuations due to wavefocusing are prominent characteristics of light in shallow marine environments. Effects of high-frequency (0.01 to 1 Hz) fluctuating light on growth rates of the red algaChondrus crispus Stackh., collected from Crane Neck Point, New York, USA, during July 1988, were determined by comparing plants grown under fluctuating and constant light regimes with similar daily irradiances. At high daily irradiance, growth rates were higher under fluctuating light than under constant light. Fluctuating light effects were frequency-dependent; growth was enhanced by fluctuations at 0.1 and 1 Hz, but not by fluctuations at 0.01 Hz. At low daily irradiance, growth rates were not affected by fluctuating light at any frequency tested. Enhancement of growth was not due to effects of high-frequency light fluctuations on photoacclimation responses ofC. crispus. Plants grown under fluctuating light at high daily irradiance actually exhibited lower photosynthetic capacity and efficiency (determined under constant light) than plants grown under constant light. These differences were attributable to variation in the density of Photosystem II reaction centers, which was low in plants grown under fluctuating light. Maximum turnover rate of whole-chain electron transport and activity of ribulose-1,5-bisphosphate carboxylase were affected by total daily irradiance, but not by high-frequency light fluctuations. Enhancement of growth under fluctuating light was partly attributable to reduced rates of dark respiration compared to rates of plants grown under constant light. The results also provided indirect evidence that high-frequency light fluctuations may enhance instantaneous photosynthetic rates. This effect could increase daily carbon gain and, therefore, stimulate growth ofC. crispus under high-frequency light fluctuations.     

Fluorescence induction in the macroalgae Chondrus crispus (Rhodophyceae) and Ulva sp. (Chlorophyceae)

The induction of in vivo chlorophyll a (chl a) fluorescence (change in fluorescence intensity during a time-scale of ms to s) was measured to determine the potential of this technique for assessing the physiological condition of the macroalgae Chondrus crispus and Ulva sp. A gradient in variable fluorescence (P-F ₀ =peak minus initial fluorescence, a measure of Photosystem II activity) was found along the frond of C. crispus, the values increasing with distance from the thallus apex. No gradient was observed for Ulva sp. thalli. Nitrogen- or phosphorus-depleted Ulva sp. required a longer dark-conditioning period and had lower values of P-F ₀ than did controls. In contrast, no differences were found in P-F ₀ of N- or P-depleted C. crispus unless values were normalized to chl a. The irradiance history of C. crispus strongly influenced P-F ₀ , even after dark-conditioning: P-F ₀ declined by about 70% as the mean daily natural irradiance increased between 2 February and 14 March 1986; a negative correlation was observed between P-F ₀ and the photon flux density 1 d prior to the measurement; P-F ₀ remained elevated when C. crispus was grown under a low photon flux density; and P-F ₀ decreased in thalli within 5 d of transfer from growth under natural irradiance to an incubator with artificial irradiance. Changes in variable fluorescence at different growth irradiances of C. crispus may be due to adaptive changes in the relative absorption cross-section of this alga. The influence of irradiance history on Ulva sp. was minimal in comparison.Issued as NRCC No. 28730Part of this study was carried out while employed by Focal Marine Ltd., Bedford, Nova Scotia, Canada     

Effect of freezing on photosynthesis of intertidal macroalgae: relative tolerance of Chondrus crispus and Mastocarpus stellatus (Rhodophyta)

The effect of freezing on photosynthetic metabolism was studied in the red algae, Chondrus crispus and Mastocarpus stellatus. Plants of both species were collected from the intertidal at Chamberlain or Kresge Point, Maine, USA (43°56N, 69°54W) between February and March 1987. Photosynthetic rates were measured immediately after freezing at-20°C and following recovery periods in seawater. Photosynthesis in C. crispus declined rapidly following freezing, falling to 70% of control values within 1 h and 30% after 3 h exposure. Minimum photosynthetic rates (7 to 9% of controls) occurred following freezing exposures of 12 h or more. Full photosynthetic recovery in C. crispus after 3 h at-20°C required 48 h. Photosynthesis in C. crispus did not fully recover in plants frozen for 6 h or more. In contrast, photosynthesis in M. stellatus was relatively unaffected by freezing exposures of <12 h. Twelve hours or more at-20°C reduced photosynthesis to 55% of controls. Photosynthesis in M. stellatus fully recovered from 24 h at-20°C within 24 h. In both species the reduction of photosynthesis by freezing was associated with damage to the plasma membrane and reduced efficiency of energy transfer from phycobilisomes to chlorophyll a, but did not appear to involve ribulose-1,5-bisphosphate carboxylase oxygenase activity. The freezing tolerance of C. crispus and M. stellatus positively correlates with their respective intertidal distributions, suggesting that freezing may be involved in controlling the distributions of these species on the shore.     

富营养化水体中菹草光合荧光特性研究

用NH4Cl、NaNO3、KH2PO4配置不同氮磷浓度的试验水,将菹草(Potamogeton cripus)分别移栽于上述水体中,观测菹草在各水体中的生长发育状况,并用水下饱和脉冲叶绿素荧光仪(Diving-PAM)测定菹草光合作用PSII有效荧光产量(Y)、光化学荧光淬灭系数(qP)、非光化学荧光淬灭系数(qN)等指标。结果表明:在TN<2mg·L-1,TP<0.4mg·L-1,Chla<118mg·m-3试验水体中,菹草长势良好,株高增加了51.2%、46.6%,叶片数增大了57.7%、58.1%;在营养盐质量浓度很高(TN8mg·L-1,TP0.8mg·L-1)、藻类生物量极高(Chla421mg·m-3)的水体中,菹草的生长受到抑制,并出现死亡。Y、qP、qN的值说明了在TN<2mg·L-1,TP<0.4mg·L-1,Chla<118mg·m-3水体中,菹草叶片吸收的光能更多的参与光化学反应,光合作用增大。高营养盐质量浓度(TN8mg·L-1,TP0.8mg·L-1,Chla421mg·m-3)水体中的菹草叶片将大部分光能用于热耗散,光合作用降低。快速光响应曲线同时也说明各水体中,菹草的最小饱和光强(Ek)都为176μmol·m-2s-1,但高营养盐质量浓度(TN8mg·L-1,TP0.8mg·L-1,Chla421mg·m-3)水体中的菹草叶片的最大光电子传递速率(rETR-max)明显低于其他水体,随着水体中营养盐浓度的升高,菹草耐光抑制能力显著降低。     

Freezing tolerance in the intertidal red algaeChondrus crispus andMastocarpus stellatus: Relative importance of acclimation and adaptation

The effect of repeated daily freezing on photosynthesis, growth and phenotypic acclimation to freezing was studied in the red algaeChondrus crispus Stackhouse andMastocarpus stellatus (Stackhouse in With.) Guiry. Algae used for experiments were collected from Chamberlain, Maine, between March and August 1987, and field observations and experiments were carried out at Chamberlain and Kresge Point, Maine between March 1987 and March 1989. After ca 30 d of daily freezing for 3 h at –5°C photosynthesis ofC. crispus was reduced to 55% of control values. Growth rates ofC. crispus were also reduced in fronds frozen daily compared to unfrozen controls, and eventually fronds became bleached and fragmented resulting in biomass losses. Fronds ofC. crispus, frozen daily, had higher photosynthetic rates following freezing events than unfrozen controls indicating that this species can acclimate to freezing conditions. Acclimation to freezing involves the light-harvesting reactions of photosynthesis. In contrast, photosynthesis and growth inM. stellatus were unaffected by repeated daily freezing for 3 h at –5°C for 36 d. No differences in photosynthesis following freezing were observed between frozen and control fronds suggesting thatM. stellatus does not phenotypically acclimate to freezing. The greater freezing tolerance ofM. stellatus relative toC. crispus results, in part, from genetic adaptations associated with plasma membranes and the light-harvesting reactions of photosynthesis.     

Carbon economy and nutrition of the alloparasitic red alga Harveyella mirabilis

Rhodomela confervoides (Huds.) Silva (Ceramiales: Rhodophyta) infected with the alloparasitic Harveyella mirabilis (Reinsch) Reinke (Cryptonemiales: Rhodophyta) was investigated with particular regard to rates of carbon fixation, patterns of soluble and insoluble assimilates, and transfer of reduced carbon from the photoautotrophic basiphyte to the fully heterotrophic epiphyte by means of ¹⁴C-labelling experiments. Gross carbon input by photosynthesis in Rh. confervoides accounted for about 53 g Cd^-1 when based on a biological unit (average individual of 200 mg fresh weight). Some 3% of the remaining net carbon gain (equivalent to about 0.7 g Cd^-1) was transferred from Rh. confervoides to its alloparasite, H. mirabilis (biological unit 0.5 mg fresh weight). Chromatographic examination showed that soluble ¹⁴C was translocated to and stored by H. mirabilis as mannosidoglycerate (=digeneaside) and probably a variety of amino acids. A certain proportion of soluble radiocarbon imported is converted by H. mirabilis to glycerolgalactoside (=floridoside). The bulk of imported ¹⁴C was recovered from the polymeric fraction, mainly from floridean starch.     

Localisation and surface quantification of secondary metabolites in the red alga Delisea pulchra

The localisation of halogenated furanones, the biologically active secondary metabolites from the red alga Delisea pulchra (Greville), was determined by a combination of fluorescence microscopy, culture studies and quantitative chemical analyses. All types of evidence showed that furanones are localised in the central vesicle of gland cells in D. pulchra. These cells release furanones onto the surface of the plant, where they can be quantified using a newly developed surface extraction-technique. Levels of furanones on the surface of the plant were highest near the apical tips (≃100 ng cm⁻²), and decreased towards the base of the alga. Variation in furanone levels within the plant and variation in the number of gland cells followed a similar pattern. The localisation of furanones within gland cells in D. pulchra and the presence and concentrations of furanones on the surface of the plant are consistent with furanones functioning as antifoulants and in mediating other ecological interactions at the surface of the alga. Received: 5 March 1998 / Accepted: 10 September 1998     

Multiple trans-Arctic passages in the red alga Phycodrys rubens: evidence from nuclear rDNA ITS sequences

In order to investigate how episodes of geological and climatic change have influenced the distribution and evolutionary diversification of Arctic to cold temperate-North Atlantic seaweed species, intraspecific genetic variation was analyzed among isolates of the sublittoral, benthic red alga Phycodrys rubens (collected between June 1992 and January 1994). Rooted phylogenetic analyses of nuclear ribosomal DNA internal transcribed spacer (ITS) sequences and the plastid encoded Rubisco spacer sequences suggest that P. rubens invaded the North Atlantic from the Pacific shortly after the opening of the Bering Strait (3 to 3.5 million years ago), colonizing both the western and eastern Atlantic coasts. Based on these data we further hypothesize that P. rubens survived along the European coasts during the more recent Pleistocene glaciations, while becoming locally extinct along the North American Atlantic coasts. Following retraction of the last ice sheet, the western Atlantic coast was colonized a second time from the Pacific. The presence of two distinct genetic types (based on ITS and Rubisco sequences) along the European coasts is postulated to be a result of isolation and subsequent differentiation. This is likely because ice-free areas are known to have existed in northern Scotland and Norway during the last glaciation. The presence of an East Atlantic genetic type along the West Atlantic coast is believed to be a recent introduction (caused by human activity) of P. rubens to Newfoundland.     

Calcium exchange and the measurement of calcification rates in the calcareous coralline red alga Amphiroa foliacea

⁴⁵Ca washout data of living thalli of Amphiroa foliacea Lamouroux in the light and dark show that there are three kinetically distinct Ca²⁺-exchanging compartments with approximate half-times (t 1/2) of 300, 20 and 3 min. The two slower compartments appear to be exchange from organic Ca²⁺-binding components of the cell wall, while the fast compartment probably represents exchange on the CaCO₃ crystal surface. Killed and decalcified thalli have a fourth compartment, with a t 1/2 of 20 to 35 min (other compartment half-times are 300, 40, 3 min), which has been identified as the greatly increased intercellular space produced during drying and decalcification. The ⁴⁵Ca and ¹⁴C uptake data show that a large proportion of the label initially taken up is into compartments other than the CaCO₃. As a result of this uptake, binding, and exchange of radioisotope, significant errors occur during the measurement of calcification rates, unless a kinetic analysis is carried out. Using such a technique, CaCO₃ calcification rates of A. folicea were measured with ⁴⁵Ca or ¹⁴C as tracers. Light stimulates calcification by up to 2.6 times, depending upon the age of the plant. Young segments have a markedly higher rate of calcification and photosythesis than do the older segments.     

Ultrastructure of tetrasporogenesis in the red alga Osmundea spectabilis var. spectabilis (Rhodomelaceae: Ceramiales: Rhodophyta)

Tetrasporogenesis in Osmundea spectabilis (Postels and Ruprecht) K.W. Nam var. spectabilis proceeds through three developmental stages. A uninucleate tetraspore mother cell with synaptonemal complexes depicted during early prophase of meiosis I comprises the youngest stage. Subsequently, the nuclear envelope obtains a highly irregular profile and is surrounded by perinuclear endoplasmic reticulum (ER) and numerous mitochondria. Proplastids with peripheral thylakoid and numerous straight-profiled dictyosomes are present. Dictyosome vesicles discharge their contents, initiating tetraspore wall formation. A four-nucleate tetraspore mother cell is formed prior to tetrahedral cleavage. During the second stage of tetraspore formation the nuclei are very irregular in outline and move toward the central region where they fuse. The formation of fibrous vacuoles originates from the fibrous vacuole associated organelles (FVAOs). Thylakoid formation from concentric lamellar bodies (CLBs) seems to occur in plastids. Extremely hypertrophied dictyosomes with two tightly compressed and densely laminated mid-regions produce vesicles which contribute to enlargement of the fibrous vacuoles. During the second stage of tetraspore development striated vesicles are formed and starch granule formation increases. Mature tetraspores are characterized by numerous cored vesicles and abundant starch granules. Plastids have a fully developed system of internal thylakoids, and cleavage is usually complete at this stage. Mature tetraspores are surrounded by a bi-layered wall and the tetrasporangial wall. Unbranched, tubular structures connecting the plasmalemma with the peripheral sheets of ER are possibly involved in wall deformation. A distinct blebbing nuclear envelope appears to sequester vesicles into the cytoplasm.     

Effect of carbon dioxide concentration on calcification in the red coralline alga Bossiella orbigniana

The relationship between various experimental concentrations of CO₂ and calcification in Bossiella orbigniana (Decaisne) was studied by measuring Ca-45 incorporation into the crystalline matrix. Air containing CO₂ at partial pressures (P_{CO 2}) of 0.04 to 5.5% was bubbled through synthetic seawater in incubation vessels. The resultant pH values in the presence of plants ranged from 6.5 to 8.7. The maximum calcification rate appears to lie between 0.11 and 1.05% P_{CO 2}. The data suggest that calcification is controlled by a biological process that may be sensitive to pH and/or to the relative bicarbonate concentration. The data also suggest that a severalfold increase in CO₂ over the present atmospheric level might lead to increased calcification in this marine alga.     

Grazer avoidance may explain the invasiveness of the red alga Gracilaria vermiculophylla in Scandinavian waters

The invasive red alga Gracilaria vermiculophylla has quickly spread across Europe, but it is unclear whether its success is based on a high tolerance to variations in environmental conditions or to the absence of native grazers that feed on this alga. We tested whether native invertebrate grazers prefer native algae to G. vermiculophylla. Feeding preferences of three common herbivores were quantified when offered G. vermiculophylla and native Fucus vesiculosus, Ceramium virgatum, and Ulva intestinalis in no-, two- and multiple-choice trials. Herbivore growth was measured when fed each of the algae separately. Grazers consumed G. vermiculophylla in no-choice trials, but avoided generally this alga when having a choice. U. intestinalis was always preferred over G. vermiculophylla, and grazers fed with U. intestinalis grew faster than those fed with G. vermiculophylla. We conclude that grazers avoid G. vermiculophylla to most native algae, which may benefit G. vermiculophylla in northern European estuaries.     

Effect of inhibitors of carbonic anhydrase activity on photosynthesis in the red alga Soliera filiformis (Gigartinales: Rhodophyta)

The effect of light intensity, pH and carbonic anhydrase (CA) inhibitors on photosynthesis of the red marine macroalgae Solieria filiformis (Kützing) Gabrielson, collected from Taliarte (Gran Canaria, Canary Islands) in 1991, has been investigated. Plants taken from the sea (wild phenotype) developed spherical morphology (ball phenotype) after 2 mo culture in aerated tanks. The photosynthetic oxygen evolution in the wild phenotype was saturated at 100 mol photons m^-2s^-1, while the ball phenotype displayed saturation at 200 mol photons m^-2s^-1. The inhibitors of total CA activity (6-ethoxizolamide) and extracellular CA activity (dextran-bound sulfonamide) inhibited photosynthesis at pH 8.2, to 90 and 50% respectively, in both phenotypes. No inhibition of the photosynthetic oxygen evolution was detected at pH 6.5. CA activity was associated with both supernatant and pellet fractions of crude extracts of S. filiformis. The rate of alkalization of the medium by the algae was dependent on light intensity. We suggest that carbon dioxide is the general form of inorganic carbon transported across the plasmamembrane in S. filiformis. HCO₃ transport into the cell takes place simultaneously by an indirect mechanism (dehydration to CO₂ catalyzed by CAext) and by direct uptake. Extracellular (CAext) and intracellular (CAint) CAs are involved in the mechanisms of inorganic carbon assimilation by S. filiformis.     

Intraspecific variation in the red alga Portieria hornemannii : monoterpene concentrations are not influenced by nitrogen or phosphorus enrichment

The red alga Portieria hornemannii (Lyngbye) Silva was selected to test the effects of enhanced nutrient availability on the production of carbon-based secondary metabolites, because of its notable site-to-site variation in monoterpene production. On Guam, the major secondary metabolite produced by this alga is ochtodene, a cyclic monoterpene. Quantitative high-performance liquid-chromatography analysis of the extracts of P. hornemannii collected from six sites on Guam showed that both ochtodene and triglyceride concentrations differed significantly among sites. Internal nitrogen and phosphorus content of the algae did not correlate with the observed variation in chemistry. Experimental enhancement of N-alone, P-alone or N + P in the field for 5 wk failed to induce a significant change in ochtodene concentrations in the alga, while triglyceride concentrations increased significantly in the N + P treatment. Ochtodene and triglyceride concentrations did not change among similar treatments in shaded (18 d) and unshaded (11 d) fertilization experiments conducted in the laboratory. Variation in ochtodene concentrations in P. hornemannii cannot be attributed to N and P availability; however, the decrease in ochtodene and triglyceride concentrations during the shaded laboratory experiment suggests that light may be a factor influencing monoterpene biosynthesis. The difference in ochtodene concentration between the initial and final sets of field controls collected for the unshaded laboratory experiment suggests that temporal variation might also contribute to differences observed among the algae at the different sites. Received 11 June 1996 / Accepted 20 July 1996     

Effect of phosphate and ammonium levels on photosynthetic and respiratory responses of the red alga Gracilaria verrucosa

Gracilaria verrucosa (Hudson) Papenfuss exposed to nutrient enriched media (0.1 mM PO₄; 1.0 mM NH ₄ ⁺ ) by pulse feeding 2 h every third day for a period of 5 wk at 20°C and 25–30 salinity showed significantly higher rates of photosynthesis regardless of photon flux density correlated with increased pigment levels. Algae in nonenriched media showed significantly higher levels of soluble carbohydrates and decreased levels of phycoerythrin and chlorophyll a. Photosynthetic and respiratory responses to temperature 15°, 25°, 30°C and salinity (15, 25, 30 S) combinations indicate broad tolerances by both nutrient enriched and non-nutrient enriched algae. Photosynthetic and respiratory rates were highest at the high temperatures. Pulse-fed algae had significantly higher photosynthetic rates than non-nutrient enriched plants at all temperature and salinity combinations. Non-nutrient enriched algae had significantly higher respiratory rates than nutrient enriched algae at only 30°C and 15. The respiratory rates of both nutrient enriched and non-nutrient algae decreased under combinations of higher temperatures and salinities. G. verrucosa, grown without nutrients, has lower tolerances to environmental stresses.     

Growth and production of the red alga Gelidium sesquipedale off the Basque coast (northern Spain)

Several aspects of the growth of the agarophyte Gelidium sesquipedale Clemente (Thuret) were studied in sublittoral populations of the Basque coast (northern Spain) from 1981 to 1986. The standing stock of the populations varied greatly with season. During two consecutive years, 60 to 70% of summer biomass had been lost by the end of the unfavourable period of year to growth (autumn and winter). The decline in biomass was mainly due to partial losses of branches, with more branches being lost from larger fronds. Yearly variations in the standing stock were also recorded. Production estimates obtained by a destructive sampling method were not satisfactory because of the spatial heterogeneity of the populations at the three locations studied. Indirect estimation of production by measuring the growth of marked fronds in a control quadrat gave more reliable estimates. Production values of 622±124, 735±147 and 615±123 g dry wt m^-2 yr^-1 were obtained at the three locations, respectively. Frond elongation rates of 7.5, 6.1, 7.5 and 7.9 cm yr^-1, respectively, were recorded during four consecutive years. In winter elongation rates of fronds were rather low. Elongation did not seem to be affected by frond length, the self-shading effect of the canopy, or changes in depth. Cuts resulting from grazing were frequent on fronds through the year, decreasing significantly with increasing depth. Sheared fronds regenerated promptly, independent of season or height at which the cut was made. For commercial purposes the best season for harvesting is summer, which is the period of maximum standing stock, infertility, and calm warm waters that provide easier working conditions for divers.