Gametogenesis and chromosome number in Postelsia palmaeformis (Laminariales, Phaeophyceae)

Gametophytes of the ‘sea palm’, the kelp Postelsia palmaeformis Ruprecht, produced gametes whether or not chelated iron was supplied in the culture medium, in contrast to the inhibition of gametogenesis seen with the absence of iron in many other kelps. As gametogenesis proceeded, every cell of the gametophytes was converted into a gamete so that the gametophytes did not continue to grow vegetatively. The portion of the life history from spore release through germination, gametophyte growth, gametogenesis, fertilization and growth of the young sporophyte was completed in 9–10 days under laboratory conditions. Chromosome counts showed that sporophytes had a diploid number of 26–34 chromosomes while sporangia and gametophytes had a haploid number of 14–17 chromosomes, indicating a typical haplodiplophasic life history as seen in other Laminariales.     

Protoplast regeneration from gametophytes and sporophytes of some species in the order Laminariales (Phaeophyceae)

Summary Protoplasts were isolated from sporophytes and from gametophyte cultures of several species in the order Laminariales. For each example, the isolation and culture procedures were investigated systematically, to identify conditions leading to plant regeneration. After dedifferentiation through a filamentous stage, protoplasts isolated from adultLaminaria saccharina sporophytes regenerated polystichous bladelets. In contrast, cells isolated fromLaminaria digitata sporophytes proved recalcitrant in culture, except when the donor plants were undifferentiated sporelings. The most critical factors for protoplast development were the origin of explants, the osmoticum used for cell isolation, cultivation in plain seawater, and the absence of stress during the first two weeks of culture. We also found that protoplast isolation from the sporophytes of members of the Laminariales results in the release of hydrogen peroxide, up to 5–120 μM final concentration in the macerating medium, a characteristic which may be related to protoplast recalcitrance. Protoplasts isolated from the gametophytic phase readily regenerated into normal gametophytes, capable of gametogenesis and producing sporophytes by fertilization.     

Development of the intercalary meristem in Chorda filum (Laminariales, Phaeophyceae) and other primitive Laminariales

Development of the intercalary meristem in the terete laminarialean species Chorda filum (L.) Stackhouse was studied in culture using light and transmission electron microscopy as well as by tracing elongation and cell divisions in various parts of the sporophyte. Growth of C. filum sporophytes could be classified into three developmental stages: (i) diffuse growth; (ii) basal meristematic growth; and (iii) intercalary meristematic growth. In the diffuse growth stage, elongation and cell division frequency were almost the same in each cell. In the basal meristematic growth stage, elongation and division of cells became localized in the tissues derived from the meristematic initial cell. Cells of the basal meristematic region contained smaller chloroplasts and many small opaque vesicles. In the intercalary meristematic growth stage, there was further elongation and differentiation of cells originating from the meristematic region, and this became more active in adjacent regions below the meristem than in regions above the meristem, causing the relative position of the intercalary meristem to shift towards the tip of the sporophyte. Meristematic cells of C. filum contained well-developed Golgi vesicles around the nucleus (perinuclear Golgi), many secretion vesicles and many small disk-shaped chloroplasts whose thylakoids were not well developed. Sporophytes of three other terete members of Laminariales, Chorda tomentosa Lyngbye, Pseudochorda nagaii (Tokida) Kawai et Kurogi, and Pseudochorda gracilis Kawai et Nabata, show diffuse growth and basal meristematic growth, but no intercalary meristematic growth. This suggests that the common ancestor of the Pseudochordaceae and Chordaceae had basal meristematic growth, and intercalary meristematic growth evolved more recently in C. filum.     

Kinetic characteristics of photoassimilate translocation in Alaria esculenta (Laminariales,Phaeophyceae)

The kinetics of translocation of ¹⁴C-labeled photoassimilate were studied in the kelp, Alaria esculenta (L.) Grev., using a Geiger-Müller detector-probe to measure radioactivity in the source and sink regions of dumbbell-shaped explants cut from blades. Rapid tracer efflux from the source occurred for 4 days following a pulse of [¹⁴C]bicarbonate, with 40–60% of the initial activity remaining in the source after 10–14 days. Portions of source and sink tissue were analysed for distribution of radioactivity in mannitol, amino-acid, organic-acid and insoluble fractions. About 75% of the radioactivity in both source and sink at the end of the experiments was in soluble organic matter. The translocation velocity of the moving solute front (1.0-1.6 cm·h^-1) was derived from time-course profiles of tracer arriving in Alaria sinks. Relative rates of translocation, calculated from these profiles, yielded skewed curves, with maximum rates of import by the sink occurring 72–96 h after the source was pulsed.     

Molecular phylogeny and timing of radiation in Lessonia (Phaeophyceae,Laminariales)

The genus Lessonia Bory de Saint‐Vincent is distributed solely in the Southern Hemisphere, with four species found in South America and five in Australasia. Our goal was to determine the evolutionary relationships between the Lessonia species of the two disjunct areas and hypothesize dispersal patterns and timing of radiation. We combined mitochondria, plastid and nuclear markers in a comprehensive dataset from multiple individuals per known species. Furthermore, for some species we added samples from multiple populations to take account of their widespread distribution over known bioregions. In all analyses the Australasian Lessonia formed a monophyletic group and in most analyses the South American species form a paraphyletic grade. Delimitations of the accepted species are highly supported except for L. variegata J. Agardh and L. nigrescens Bory de Saint‐Vincent. We showed for the first time four lineages for the New Zealand endemic Lessonia variegata with an unexpected high level of genetic differentiation. Our analysis cannot resolve whether the dispersal of the common ancestor of this genus was from the Americas to Australasia or vice versa. Molecular clock analysis suggested that a sudden radiation took place in Australasia c. 3.5 Mya when almost all Australasian species diverged within a time frame of only 35 000 years.     

Temperature requirements for the growth of young sporophytes of Undaria pinnatifida and Undaria undarioides (Laminariales, Phaeophyceae)

The relative growth rate of young sporophytes of Undaria pinnatifida (Harvey) Suringar and Undaria undarioides (Yendo) Okamura was examined in order to understand the difference in distribution of these two species around the coast of Japan. The optimal temperature for growth of both species was similar at 20°C and the upper critical temperature for growth was also similar, at 27°C for U. pinnatifida and 26°C for U. undarioides. Therefore, the optimal and upper critical temperatures for growth of the young sporophytes are not the main factors determining the distribution of each species. Next, the lower critical temperatures for growth were examined. For the young sporophytes of U. pinnatifida, the lower limit was less than 5°C while for those of U. undarioides it was 15°C. Thus, the difference in the lower critical temperature for growth between the two species was approximately 10°C. During the period of young sporophyte growth in the field, the temperature at the mouth of Ise Bay, Japan, where U. pinnatifida occurs, ranges from 12.7°C in December to 13.1°C in April, with a minimum of 7.9°C in February. Our experiments indicate that young sporophytes are able to grow throughout this period. The temperature off Hamajima, Japan, where U. undarioides occurs, ranges from 19.1°C to 14.8°C during the same time period. Again, young sporophytes are able to growth throughout this period, although minimum winter temperatures are only just high enough for growth. These natural temperature ranges during the growth season of the sporophytes agree well with the experimentally determined temperature requirements for growth of each species. Therefore, the difference between the two species in the critical temperature required for growth of the young sporophytes, especially in the low temperature range, is one of the major factors determining the distribution pattern of each species.     

Taxonomy of the southwestern Atlantic endemic kelp: Laminaria abyssalis and Laminaria brasiliensis (Phaeophyceae,Laminariales) are not different species

Two endemic species of Laminaria, Laminaria abyssalis Joly & Oliveira Filho and L. brasiliensis Joly & Oliveira Filho, from the tropical southwestern Atlantic coast have been described. The aim of this work was to determine the conspecificity of these species based on morphological and molecular analyses (ribulose 1,5‐bisphosphate carboxylase/oxgenase, large subunit (rbcL), internal transcribed spacer (ITS) and cytochrome c oxidase subunit I (coxI)). We found an overlap between the morphological characters that are considered taxonomically important for distinguishing these two species; these characters included a differing pattern of blade splitting. In the three molecular analyses, the Brazilian Laminaria specimens were grouped into one clade with maximum support. These data support the hypothesis that the individuals analyzed represent only one species, L. abyssalis. The molecular analysis also showed L. abyssalis to be sister group to L. digitata.     

Compensatory growth of the kelp Macrocystis integrifolia (Phaeophyceae, Laminariales) against grazing of Peramphithoe femorata (Amphipoda, Ampithoidae) in northern-central Chile

Compensation of tissue loss has been considered an alternative strategy for seaweeds that have no or only minor chemical or structural defense against herbivory. Compensatory responses are facilitated by resource transfer among different tissues and have been suggested for large kelps. Macrocystis integrifolia (Bory) is a common kelp species from northern-central Chile, which is characterized by high growth rates and the absence of lipophilic chemical defenses against herbivore grazing. Herein, we used the giant kelp M. integrifolia to test for compensatory growth in response to grazing by the nest-dwelling amphipod Peramphithoe femorata (Krøyer). Amphipods were allowed to graze inside nests on subapical blades of M. integrifolia sporophytes for 14 days. We measured growth and chemical composition (C, N, laminaran and mannitol) of apical and subapical blades of grazed and ungrazed (control) sporophytes. Our results revealed the capability of M. integrifolia to maintain elongation rates in grazed subapical blades, which were similar to those of subapical blades from ungrazed sporophytes. Apical blades grew slower in grazed than in ungrazed sporophytes indicating a trade-off between apical and subapical blades when herbivores were present. Thus, compensation occurs in blades directly attacked by grazers and is probably mediated by vertical resource allocation within sporophytes to subapical blades, a suggestion supported by the fact that stipe internodes in these regions grew more on grazed sporophytes. In general, our study indicates that M. integrifolia exhibits compensatory growth against the herbivore amphipod P. femorata, and we suggest that this could be an important strategy of large kelp species to tolerate moderate grazing intensities.     

Species delimitation and phylogeographic analyses in the Ectocarpus subgroup siliculosi (Ectocarpales,Phaeophyceae)

The genus Ectocarpus (Ectocarpales, Phaeophyceae) contains filamentous algae widely distributed in marine and estuarine habitats of temperate regions in both hemispheres. While E. siliculosus has become a model organism for genomics and genetics of the brown macroalgae, accurate species delineation, distribution patterns and diversity for the genus Ectocarpus remain problematic. In this study, we used three independent species delimitation approaches to generate a robust species hypothesis for 729 Ectocarpus specimens collected mainly along the European and Chilean coasts. These approaches comprised phylogenetic reconstructions and two bioinformatics tools developed to objectively define species boundaries (General Mixed Yule Coalescence Method and Automatic Barcode Gap Discovery). Our analyses were based on DNA sequences of two loci: the mitochondrial cytochrome oxidase subunit 1 and the nuclear internal transcribed spacer 1 of the ribosomal DNA. Our analyses showed the presence of at least 15 cryptic species and suggest the existence of incomplete lineage sorting or introgression between five of them. These results suggested the possible existence of different levels of reproductive barriers within this species complex. We also detected differences among species in their phylogeographic patterns, range and depth distributions, which may suggest different biogeographic histories (e.g., endemic species or recent introductions).     

Temperature requirements for the growth and maturation of the gametophytes of Undaria pinnatifida and U. undarioides (Laminariales, Phaeophyceae)

Gametophytes of two Undaria species, U. pinnatifida and U. undarioides (Laminariales, Phaeophyceae), were studied to determine their water temperature requirements in order to understand their different distributions in Mie Prefecture, Japan. The optimal temperature for growth was 20°C for gametophytes of both species, and the upper critical temperature for growth was also the same for both species at 28°C. Therefore, the optimal and critical temperatures for growth of the gametophytes are not the main factors determining distribution. The optimal temperature for maturation of U. pinnatifida was approximately 10–15°C, whereas it was closer to 20–21°C for U. undarioides, a difference between these species of at least 5°C. In autumn and early winter, the seawater temperature at the mouth of Ise Bay, where U. pinnatifida is distributed, ranges from 21.6°C (October) to 12.7°C (December), and off Hamajima, where U. undarioides is found, the range is from 22.7°C (October) to 19.1°C (December). The seawater temperatures from October to December, which is the maturation season for the gametophytes, agreed well with the optimal temperature requirements for maturation of the gametophytes of both species. Thus the difference in the maturation temperature range of the gametophytes is a major factor determining distribution of these Undaria species along the Japanese coast.     

Sporogenic and vegetative tissues of Saccharina latissima (Laminariales,Phaeophyceae) exhibit distinctive sensitivity to experimentally enhanced ultraviolet radiation : photosynthetically active radiation ratio

Fertile Saccharina latissima sporophytes, collected in the Kongsfjorden, Ny‐Ålesund, Spitsbergen, Norway (78°56.87′ N, 11°51.64′ E) were investigated in relation to its sensitivity to experimentally enhanced ultraviolet radiation : photosynthetically active radiation (UVR : PAR) ratios. Irradiance of UVR were 4.30 W m^?2 of UV‐A (320–400 nm) and 0.40 W m^?2 of UV‐B (280–320 nm), and PAR (400–700 nm) was ～4.30 W m^?2 (=20 µmol photons m^?2 s^?1). Excised soral (sporogenic) and non‐soral (vegetative) tissues were separately irradiated for 16 h at 7°C. Transmission electron microscopy showed abundant occurrence of physodes, electron dense particles (～300–600 nm) in the sorus. Paraphysis cells, with partly crystalline content, large mitochondria and abundant golgi bodies were towering over the sporangia. In soral tissue, cells were not visibly altered by the PAR + UVR irradiation. The chloroplasts, flagella and nucleus of unreleased meiospores inside the sporangial parent cells were visibly intact. Severe changes in the chloroplast structure of vegetative tissue occurred after PAR + UVR irradiation. These changes included wrinkling and dilatation of the thylakoid membranes, and appearance of electron translucent areas inside the chloroplasts. In vegetative cells exposed to PAR + UVR, the total amount of physodes, was slightly higher as in cells exposed to PAR only. Initial values of optimum quantum yield of photosystem II (F_v/F_m) were 0.743 ± 0.04 in non‐soral and 0.633 ± 0.04 in soral tissue. Vegetative tissue was observed to be more sensitive to radiant exposure of PAR and PAR + UVR compared to reproductive tissue. Under PAR, a 20% reduction in Fv/Fm was observed in non‐soral compared to no reduction in soral tissue, whereas under PAR + UVR, 60% and 33% reduction in Fv/Fm was observed in non‐soral and soral tissues, respectively. This can be attributed to the corresponding three times higher antiradical power (ARP) capacity in soral compared to non‐soral tissue.     

Effects of nutrient enrichment and herbivory on morphology,reproduction and chemical content of Turbinaria conoides (Phaeophyceae)

The effects of nutrient enrichment and herbivory on resource allocation patterns among morphology, reproduction, and chemical content of the brown alga, Turbinaria conoides (J. Agardh) Kützing were tested in the shallow subtidal zone of the Gulf of Thailand. The field experimental design comprised 36 plots (50 × 50 cm²) with and without herbivores, and two nutrient levels. Cages (uncaged and fully caged plots) were used to exclude herbivorous fishes and two nutrients levels were achieved by experimental enrichment above ambient nutrient concentrations. For morphology of Turbinaria, the maximum length (holdfast to frond apex), the width of the base of the stipe, number of blades, holdfast and stipe diameter were measured. Biomass, reproductive output, and tissue nutrient (carbon: nitrogen: phosphorus) content of T. conoides were examined. Phlorotannin concentrations were examined using a modified Folin‐Ciocalteu method. The results showed that herbivory had no effect on morphology, reproduction, or phlorotannin concentrations. This could be due to the structural and morphological deterrents of alga, which might minimize grazing effects from herbivorous fishes. Nutrient enrichment had no effect on morphology and reproduction of T. conoides, possibly due to low nutrient demand in Turbinaria. However, nutrient enrichment did affect phlorotannin concentrations, as phlorotannins in the enriched plots were lower than the ambient plots. These results support, in part, the carbon–nutrient balance hypothesis that states algae will allocate fewer resources to production of anti‐herbivore chemical compounds when enriched with potential growth‐limiting nutrients.     

The edible brown seaweed Alaria esculenta (Phaeophyceae, Laminariales): hybridization, growth and genetic comparisons of six Irish populations

Alaria esculenta populations from six differentgeographical locations on the Irish coast wereexamined for hybridization abilities, growth rates andgenetic make-up with a view towards identifying afast-growing strain suitable for aquaculture.Hybridization experiments under laboratory conditionswith the three most geographically dispersedpopulations showed that all cross combinations wereinterfertile, although differences were found insurvival, and in blade and hapteron morphology. Acomparison of relative growth rates showed significantdifferences amongst the self-crosses and hybrids. Thedata of the hybridization experiments and growth ratesunder laboratory conditions show that the bestpopulation for the purpose of seaweed aquaculture arethe Slea Head and Corbet Head self-crosses and theirhybrids. Genetic fingerprinting of the internaltranscribed spacer of the ribosomal DNA of five A. esculenta isolates from geographically separatedpopulations in Ireland revealed no restriction lengthpolymorphisms between the tested isolates and showthat the A. esculenta populations around theIrish coast are clearly genetically homogenous inrespect of the DNA region examined. The geneticanalysis, interfertility of the populations,morphology and growth rates are discussed with a viewto potential cultivation.     

Revision of the kelp family Alariaceae and the taxonomic affinities of Lessoniopsis Reinke (Laminariales,Phaeophyta)

The morphologically diverse members of the Laminariales are separated into three families, the Alariaceae, Laminariaceae and Lessoniaceae, on the basis of developmental features exhibited at the stipeblade transition zone. We have investigated the relationships of lessoniacean taxa with those of alariacean and laminariacean affinity. This was done by completing phylogenetic analyses of 3 small-subunit (SSU) rRNA gene (113 bp), first internal transcribed spacer (ITS 1), 5.8S rRNA gene and second internal transcribed spacer (ITS2)(12 bp) sequence data from a variety of taxa. In summary, the Alariaceae and Lessoniaceae are polyphyletic. We present a restricted view of the Alariaceae, including only Alaria and Pterygophora of the genera usually placed in this family, and transfer the monotypic genus Lessoniopsis to this group. Current theories of kelp evolution and biogeography must be reconsidered in view of our data.     

Streblonema (Ectocarpales,Phaeophyceae) infection in the kelp Laminaria saccharina (Laminariales,Phaeophyceae) in the western Baltic

The brown alga Laminaria saccharina is the dominant subtidal macroalga in Kiel Bay, western Baltic. It is infected by the microscopic brown alga, Streblonema aecidioides. Infected thalli may show symptoms of Streblonema disease, i.e. alterations of blade and stipe, ranging from dark spots to heavy deformations and completely crippled thalli. Samples taken from a single locality all year round show that (i) the host population is infected at a high rate of 87±13% (SD), but that (ii) a considerable proportion of thalli containing Streblonema does not show disease symptoms, and that (iii) juvenile hosts, which mainly appear in autumn, are infected at almost the same rate. Thus the infection seems to occur early in the host's life. Juveniles in nature show fewer symptoms of the disease than adults. Two months after infection, oxygen production and growth in laboratory-raised experimentally infected juvenile hosts was not different from uninfected controls. Experimental thalli showed more severe morphological alterations than uninfected controls only four months after infection. Both field and laboratory observations indicate that a lag phase exists between infection and outbreak of the disease.     

Phylogeography of Padina boryana (Dictyotales,Phaeophyceae) around the Thai-Malay Peninsula

The Indo-Pacific Ocean is a biodiversity hotspot for marine organisms. In this area, most of the research has focused on marine animals, such as reef fish, molluscs and other associated coral fauna, but very little has been done on macroalgae. The Thai-Malay Peninsula is an important north–south barrier in this area, which faces two different oceans – the Indian Ocean and the Pacific Ocean. This study aims to investigate genetic distribution patterns of Padina boryana Thivy around the Thai-Malay Peninsula, where it is common. Three DNA marker regions, the mitochondrion-encoded cytochrome c oxidase subunit 3 gene (cox3); the plastid rbcL, and the nuclear internal transcribed spacer 2 (ITS2) were used to evaluate genetic diversity and the relationships within and between populations. Samples were collected from both the Andaman Sea and Gulf of Thailand sides of the peninsula. Parsimony networks and maximum likelihood and Bayesian analyses showed clearly that there are two separated P. boryana lineages, one restricted to the Gulf of Thailand and the other to the Andaman Sea and other areas of the Indo-Pacific. The effect of different ocean currents along the Andaman Sea and Gulf of Thailand may have shaped these populations of P. boryana. This phylogeographic separation, based on persistent currents in the area, may affect other marine organisms along the Thai peninsula.     

Oligocarrageenans and tissue-dependant oxidative burst in Solieria chordalis (Rhodophyceae, Gigartinales)

The release of hydrogen peroxide by thallus fragments of the rhodophycean Solieria chordalis (C. Agardh) J. Agardh has been documented both in the presence and in the absence of oligosaccharides. Within 1 h, ramuli were able to release large amounts of peroxide in the absence of any chemical stress. Among potential elicitors tested, only degree of polymerization 1 (DP1) and DP7‐8 oligo‐iota‐carrageenans stimulated defense mechanisms in both axes and ramuli as shown by the occurrence of an oxidative burst. Chopping of the tissues had no effect on the intensity of the burst, therefore suggesting that mainly cortical cell layers were involved in the process. After 5 min incubation, a dose of 125 μg mL^?1 of an oligomeric mixture containing a large proportion of DP1 units proved to be sufficient to obtain a maximal response. The intensity of the burst was significantly higher with isolated ramuli than with pieces of the axis, with outer peroxide accumulations reaching 200 nmol g^?1 fresh weight of treated tissue. Altogether, our results show that S. chordalis is able to react to a simulated pathogen attack by an oxidative burst and that the capacity to carry out an oxidative burst is stronger in ramuli than in axes.     

Desmarestia antarctica (Desmarestiales,Phaeophyceae), a new ligulate Antarctic species with an endophytic gametophyte

Desmarestia antarctica Moe & Silva, a new species from the Antarctic Peninsula and the Scotia Arc, is distinguished from all other ligulate members of the genus by the arrangement of zoosporangia, which are produced together with paraphyses in a raised sorus. Among all species in the genus, onlyD. anceps Montagne shares this feature.Desmarestia antarctica is the first species of the genus for which an endophytic gametophyte is demonstrated, the usual host beingCurdiea racovitzae De Wildeman, a red alga of the familyGracilariaceae. The sporophyte contains only a moderate concentration of acid compared to Northern Hemisphere ligulate species, while physodes that probably contain polyphenolic compounds form noticeable speckles and dark margins in preserved plants. Often more than one axis arises from a single holdfast, probably as the result of compounding rather than proliferation.Dedicated to Prof. DrLothar Geitler on the occasion of the 90th anniversary of his birthday.     

Sequence repeats enlarge the internal transcribed spacer 1 region of the brown alga Colpomenia sinuosa (Scytosiphonaceae, Phaeophyceae)

Colpomenia sinuosa is an annual brown algal species that occurs in temperate to tropical waters of the world. In order to examine the genetic diversity among populations of the species and to discuss its current distribution, we analyzed the nuclear ribosomal DNA internal transcribed spacer (ITS) region from 18 specimens of C. sinuosa , and, for comparison, plastid rbc L from the same specimens. The C. sinuosa ITS region (2141–2534 base pairs) is approximately 2.2 times the length of most other brown algae. We found a long repeated sequence of approximately 190 base pairs in the first half of the ITS1 region; five repeats in Northern Hemisphere collections and three in those from the Southern Hemisphere, which result in ITS length variation. The unequal occurrence of tandem repeats of C. sinuosa corresponds to the geographical distribution of the species. The rbc L sequences from all the specimens of C. sinuosa , except the Canary Island samples, were identical, indicating that they indeed belong to the same species.     

Dark respiration of the stipe of Ecklonia cava (Laminariales, Phaeophyta) in relation to temperature

Sporophytes of Ecklonia cava Kjellman (Laminariales, Phaeophyta) with a stipe length of 22–102 cm were collected at 6–9 m depth in Nabeta Bay, Shimoda, central Japan by scuba diving in February (winter) and in August (summer) 1998. Dark respiration of the intact stipe of E. cava was measured at various water temperatures ranging from 15 to 27.5°C in winter and 15–30°C in summer in a closed system by using a dissolved oxygen meter. The stipe respiration was compared on whole stipe, length, surface area, volume, wet weight and dry weight bases. On each basis, the stipe respiration always increased with a rise in water temperature within the temperature range investigated. The stipes showed similar respiration rates on each basis of length, surface area, volume, wet weight and dry weight at each temperature, irrespective of the stipe length. The mean respiration rates in winter (at 15–27.5°C) were: length, 16.7–32.5 μL O₂ cm^?1 h^?1; surface area, 3.2–6.2 μL O₂ cm^?2 h^?1; volume, 7.6–15.0 μL O₂ cm^?3 h^?1; wet weight, 6.2–12.2 μL O₂ g (wet weight)^?1 h^?1; and dry weight, 43.8–88.0 μL O₂ g (dry weight)^?1 h^?1. Those for summer (at 15–30°C) were: length, 17.1–32.0 μL O₂ cm^?1 h^?1; surface area, 3.6–6.8 μL O₂ cm^?2 h^?1; volume, 9.7–18.7 μL O₂ cm^?3 h^?1; wet weight, 7.6–14.6 μL O₂ g (wet weight)^?1 h^?1; and dry weight, 49.4–95.8 μL O₂ g (dry weight)^?1 h^?1. This is the first report of the intact stipe respiration of E. cava at various temperatures.