GENETICS OF CHAMPIA PARVULA (RHODYMENIALES. RHODOPMYTA): MENDELIAN INHERITANCE OF SPONTANEOUS MUTANTS1,2

Spontaneous mutants affecting the morphology and pigmentation of the small marine red alga Champia parvula were genetically characterized. All of the variants were transmitted in the Mendelian pattern expected for recessive nuclear mutations. The results permit the identification of four mutant genes affecting morphology and four mutant genes affecting pigmentation. One of the latter, designated yell, was interesting in that it also appeared to act as a recessive lethal during carposporophyte development. So linkage bas been discovered in crosses conducted thus far. Champia parvula was found to have many desirable features for genetic studies, especially a very rapid sexual life cycle, and it is suggested that this species should be considered as a possible model organism for future genetic studies on the Rhodophyta or for teaching seaweed genetics.     

CELL DIVISION RATES OF EUCARYOTIC ALGAE MEASURED BY TRITIATED THYMIDINE INCORPORATION INTO DNA: COINCIDENT MEASUREMENTS OF PHOTOSYNTHESIS AND CELL DIVISION OF INDIVIDUAL SPECIES OF PHYTOPLANKTON ISOLATED FROM NATURAL POPULATIONS1

The cell cycle marker event of DNA replication in eucaryotic algae was identified using ³H-Thymidine (³H-TdR) incorporation. The frequency of cells (F) within a population undergoing DNA replication was estimated and the cell division rate (μF) calculated. In laboratory cultures the rates of cell division calculated from changes in cell numbers (μN) and μF were similar. Dual labelling with ³H-TdR and NaH¹⁴CO₃ enabled rates of cell division and photosynthesis to be coincidently measured for individual species of algae. Using these single species radioisotope techniques, several distinct photosynthesis irradiance and cell division irradiance relationships were found for: (1) different species of phytoplankton isolated from the same sample, and (2) the same species isolated from different environments. These techniques allow the coupling between photosynthesis and cell division to be examined with high resolution for algae in situ.     

PRODUCTIVITY OF THE FILAMENTOUS ALGA PITHOPHORA OEDOGONIA (CHLOROPHYTA) IN SURREY LAKE,INDIANA1

Biomass, akinete numbers, net photosynthesis, and respiration of Pithophora oedogonia were monitored over two growing seasons in shallow Surrey Lake, Indiana. Low rates of photosynthesis occurred from late fall to early spring and increased to maximum levels in late spring to summer (29–39 mgO₂·g^?1 dry wt·h^?1). Areal biomass increased following the rise in photosynthesis and peaked in autumn (163–206g dry wt·m^?2). Photosynthetic rates were directly correlated with temperature, nitrogen, and phosphorus over the entire annual cycle and during the growing season. Differences in photosynthetic activity and biomass between the two growing seasons (1980 and 1981) were apparently related to higher, early spring temperatures and higher levels of NO₃-N and PO₄-P in 1981. Laboratory investigations of temperature and light effects on Pithophora photosynthesis and respiration indicated that these processes were severely inhibited below 15°C. The highest P_max value occurred at 35°C (0.602 μmol O₂·mg^?1 chl a·min^?1). Rates of dark respiration did not increase above 25°C thus contributing to a favorable balance of photosynthetic production to respiratory utilization at high temperatures. Light was most efficiently utilized at 15°C as indicated by minimum values of I_k(47 μE·m^?2·s^?1) and I_c (6 μE·m^?2·s^?1). Comparison of P. oedogonia and Cladophora glomerata indicated that the former was more tolerant of temperatures above 30°C. Pithophora's tolerance of high temperature and efficient use of low light intensity appear to be adaptive to conditions found within the dense, floating algal mats and the shallow littoral areas inhabited by this filamentous alga.     

Responses of crustose corallines to epiphyte and canopy cover

Although a dense cover of epiphytes is generally considered to be harmful for some coralline algae (Corallinaceae, Rhodophyta), crustose corallines in the littoral zone seem to be preserved from bleaching when covered by canopy plants and epiphytes during summer. This study aimed to test the responses of coralline crusts to epiphytes and canopy algae and their interaction with grazing limpets. Growth rates and color changes were followed in two crust species in areas with or without canopy algae in the Isle of Man, British Isles. Limpets were excluded, to allow epiphytes to grow upon crusts. Responses were measured both on pieces of crusts upon acrylic plates and on crusts growing naturally on the shore. Fucus canopy and epiphytic Enteromorpha significantly influenced the crusts' growth, depending on season. Epiphytes reduced the light levels beneath by up to 78%, more than the canopy algae (62%). Crusts exposed outside the canopy bleached in summer, but gradually restored their color once they were covered by epiphytes. The fast-growing Phymatolithon lenormandii (Aresch.) Adey recovered its coloration more quickly than the slow-growing P. purpureum (P. et H. Crouan) Woelkerling et Irvine. However, neither crust species could restore its color when epiphytes were reduced by grazing limpets, Patella vulgata L. Bleaching did not kill the crusts, but seemed to interfere with crusts' growth. Restoration of pigmentation was quantified for the first time on bleached coralline crusts. Epiphyte and canopy algae were experimentally shown to be beneficial, probably by providing shade and also protecting crusts from desiccation.     

Photosynthetic light-response curves and photoinhibition of the deep-water laminaria abyssalis and the intertidal laminaria digitata (phaeophyceae)

Photosynthetic light‐response curves of the deep‐water Laminaria abyssalis Oliveira and of the intertidal L. digitata Lamoroux were determined and related to photoinhibition phenomena as monitored by oxygen evolution and photosystem II efficiency (F_V/F_M). L. abyssalis has half the pigment content, number of cells and plastids, and photosynthetic capacity per unit area compared with L. digitata. L. abyssalis showed a higher in vivo Chl a absorption coefficient and higher photosynthetic efficiency on a Chl a basis, although the two algae showed somewhat similar light‐response curves on a Chl a basis. Both species showed similar Chl a/Chl c and Chl a/fucoxanthin ratios, and similar dark respiration rates and light compensation points. In addition, they also showed similar convexities in their light‐response curves and no differences in their light saturation of F_V/F_M. Room temperature chlorophyll fluorescence induction measurements of fronds incubated in 3‐(3,4‐dichlorophenyl)‐1,1‐dimethylurea (DCMU) suggest that both species may have a similar PSII absorption cross section. Thus, L. abyssalis appears to optimize its light absorption at very low light intensities, not by increasing the pigment content, but by absorbing light more efficiently. However, L. abyssalis was more sensitive to photoinhibition than L. digitata and showed no recovery of F_V/F_M and O₂ evolution after a photoinhibitory treatment, even with a subsequent exposure to 24 h of dim light. L. digitata, on the other hand, recovered its photosynthetic capacity within 6 h under dim light. These results suggest that photosynthetic light‐induction curves based on Chl a are not a good indicator of either the photosynthetic capacity or the sensitivity to photoinhibition when macroalgae of different species are being compared. Based on their light‐response and photoinhibition characteristics, we suggest that L. abyssalis, a deep‐water oceanic macroalgae, is an atypical shade alga whereas L. digitata has the properties of a sun alga.     

TAXONOMY AND PHYLOGENY OF OSMUNDEA (RHODOMELACEAE, RHODOPHYTA) IN ATLANTIC EUROPE

Two species of Osmundea Stackhouse (Rhodomelaceae, Rhodophyta) that occur in Atlantic Europe have been confused under the names Osmundea ramosissima (Oeder) Athanasiadis and Osmundea truncata (Kützing) Nam et Maggs, regarded until now as a synonym of O. ramosissima. An epitype from its type locality (Stavanger, Norway) is selected for Osmundea ramosissima Athanasiadis, recognized here as a valid name for Fucus ramosissimus Oeder, nom. illeg. Details of vegetative and reproductive morphology of O. ramosissima are reported, based on material from France, the British Isles, and Helgoland. Osmundea ramosissima resembles other species of Osmundea in its vegetative axial segments with two pericentral cells and one trichoblast, spermatangial development from apical and epidermal cells (filament type), the formation of five pericentral cells in the procarp‐bearing segment of the female trichoblast, and tetrasporangial production from random epidermal cells. Among the species of Osmundea, O. ramosissima is most similar to O. truncata. Both species have discoid holdfasts, secondary pit connections between epidermal cells, and cup‐shaped spermatangial pits. They differ in that: (a) O. ramosissima lacks lenticular wall thickenings and refractive needle‐like inclusions in medullary cells, both of which are present in O. truncata; (b) O. ramosissima has branched spermatangial filaments that terminate in a cluster of several cells, whereas in O. truncata the unbranched spermatangial filaments have a single large terminal sterile cell; and (c) cystocarps of O. ramosissima lack protuberant ostioles but ostioles are remarkably protuberant in O. truncata. Phylogenetic analyses of rbcL sequences of Laurencia obtusa (Hudson) Lamouroux and all five Atlantic European species of Osmundea, including the type species, strongly support the generic status of Osmundea. Osmundea ramosissima and O. truncata are closely related (5.2% sequence divergence) and form a well‐supported clade sister to a clade consisting of O. pinnatifida (Hudson) Stackhouse, O. osmunda Stackhouse and O. hybrida (A. P. de Candolle) Nam. The formation of secondary pit connections between epidermal cells is a synapomorphy for the O. ramosissima+O. truncata clade. The close relationship between species with cup‐shaped spermatangial pits (Osmundea hybrida) and urn‐shaped pits (Osmundea pinnatifida and Osmundea osmunda) shows that spermatangial pit shape is not an important phylogenetic character. Parsimony analysis of a morphological data set also supports the genus Osmundea but conflicts with the molecular trees in infrageneric relationships, placing O. hybrida basal within the Osmundea clade and grouping O. osmunda and O. pinnatifida but not O. truncata and O. ramosissima. A key to Osmundea species is presented.     

Studies in the Liagoraceae (Nemaliales, Rhodophyta) I. The genus Trichogloea

The calcified, though strongly mucosoid gametophytes of Trichogloea (Nemaliales, Liagoraceae) usually appear seasonally for a short period in tropical and subtropical regions. Vegetative and reproductive characteristics of the presently recognized species, Trichogloea requienii (Mont.) Kützing (the generitype), Trichogloea lubrica J. Agardh, and Trichogloea herveyi W. R. Taylor, are described and compared. In addition, the identities of two previously synonymized species, Trichogloea jadinii Børgesen and Trichogloea javensis B0rgesen are clarified. Species of the genus display considerable variation in habit and branching patterns, necessitating an assessment of reproductive structures for accurate identification. Reliable features of Trichogloea include the structure of the medulla and the cortical (assimilatory) filaments, location and structure of carpogonial branches and spermatangia, and the nature of sterile filaments in the vicinity of the cystocarp. Secondary features include both external and internal branching pattern, and the proportion of calcium carbonate to lubricous material, the latter having to be assessed from fresh collections. The present study provides a comparative morphological account of the species of Trichogloea. As a result, T. requienii, T. lubrica, T. herveyi are confirmed as independent species; T. jadinii is removed from synonymy with T. lubrica and placed in that of T. requienii, and T. javensis is confirmed as a synonym of Izziella orientalis (J. Agardh) Huisman et Schils.     

Low algal carbon content and its effect on the C : P stoichiometry of periphyton

1. We examined the contribution of algal cells to periphytic organic carbon and assessed the effects of variable biomass composition on the carbon : phosphorus (C : P) ratio of periphyton. We compiled more than 5000 published and unpublished observations of periphytic carbon : chlorophyll a (C : Chl) ratios, an index of algal prevalence, from a variety of substrata collected from lake and low‐salinity coastal habitats. In addition, we converted estimates of algal biovolume into algal C to obtain an independent measure of cellular algal carbon in periphyton. This information was used in a model relating periphyton C : P ratio to algal cellular carbon, the algal C : P ratio, and the C : P ratio of non‐algal organic matter in periphyton. 2. The mean C : Chl ratio of periphyton (405) was relatively high with values in >25% of the samples exceeding 500. On average, 8.4% of total periphyton C was accounted for by C in algal cells. Only 15% of samples were found to have more than 15% periphyton C in cellular algal carbon. Our model showed a nonlinear relationship between periphytic C : P ratios and the C : P ratio of algal cells in the periphyton when non‐algal organic matter was present. However, even at relatively low cellular algal C (<10% of total C), algal C : P ratios can strongly affect the C : P ratio of periphyton as a whole (i.e. algal cells plus other organic matter). 3. The high C : Chl ratios and the low biovolume‐derived algal C of periphyton samples in our data set indicate that algal cells are typically a minor component of organic carbon in periphyton, However, this minor contribution would not preclude algal cellular stoichiometry from notably influencing periphyton C : P ratios.     

Benthic–pelagic coupling in the population dynamics of the harmful cyanobacterium Microcystis

1. In eutrophic lakes, large amounts of the cyanobacterium Microcystis may overwinter in the sediment and re‐inoculate the water column in spring. 2. We monitored changes in pelagic and benthic populations of Microcystis in Lake Volkerak, The Netherlands. In addition, sedimentation rates and the rate of recruitment from the sediment were measured using traps. These data were used to model the coupling between the benthic and pelagic populations and to calculate the contribution of overwintering benthic and pelagic populations to the magnitude of the pelagic summer bloom. 3. Changes in the benthic Microcystis population showed a time lag of 3–14 weeks compared with the pelagic population. This time lag increased with lake depth. The largest amount of benthic Microcystis was found in the deepest parts of the lake. These observations suggest horizontal transport of sedimented Microcystis from shallow to deep parts of the lake. 4. Recruitment from and sedimentation to the sediment occurred throughout the year, with highest recruitment and sedimentation rates during summer. Model simulations indicate that the absence of benthic recruitment would reduce the summer bloom by 50%. 5. In spring, the total pelagic population was three to six times smaller than the total benthic population. Yet, model simulations predict that the absence of this small overwintering pelagic population would reduce the summer bloom by more than 64%. 6. Reduction of the overwintering pelagic populations, for instance by flushing, may be a useful management strategy to suppress or at least delay summer blooms of Microcystis.     

Fitness effects of new mutations in Chlamydomonas reinhardtii across two stress gradients

Most spontaneous mutations affecting fitness are likely to be deleterious, but the strength of selection acting on them might be impacted by environmental stress. Such stress‐dependent selection could expose hidden genetic variation, which in turn might increase the adaptive potential of stressed populations. On the other hand, this variation might represent a genetic load and thus lead to population extinction under stress. Previous studies to determine the link between stress and mutational effects on fitness, however, have produced inconsistent results. Here, we determined the net change in fitness in 29 genotypes of the green algae Chlamydomonas reinhardtii that accumulated mutations in the near absence of selection for approximately 1000 generations across two stress gradients, increasing NaCl and decreasing phosphate. We found mutational effects to be magnified under extremely stressful conditions, but such effects were specific both to the type of stress and to the genetic background. The detection of stress‐dependent fitness effects of mutations depended on accurately scaling relative fitness measures by generation times, thus offering an explanation for the inconsistencies among previous studies.