GROWTH AND PHOTOSYNTHESIS OF ULVA ROTUNDATA (CHLOROPHYTA) AS A FUNCTION OF TEMPERATURE AND SQUARE WAVE IRRADIANCE IN INDOOR CULTURE1

Temperature and photon flux density (PFD) vary independently in estuaries, e.g. high PFD may occur at any temperature, so it is necessary to consider synergistic effects of these factors on algal growth. Because natural PFD is highly variable and daylength changes confound seasonal temperature cycles, it is easier to interpret factorial experiments in controlled laboratory conditions. Clonal Ulva rotundata Blid. (Chlorophyta) has been studied extensively in outdoor culture. In this study it was maintained indoors under square wave photoperiods at five PFDs and three temperatures. Growth rate, photqsynthetic light response (P-I) curves, and photosystem II chlorophyll fluorescence properties were measured at the growth temperature following acclimation. Interactions between PFD and growth temperature were strongly indicated in all physiological parameters measured. Greatest PFD response occurred at the highest temperature, and the largest temperature response occurred at the highest PFD. Light-saturated photosynthesis (P_m) dark respiration (R_d), and light-limited quantum yield (Φ_m) were sufficient to describe acclimation status. The light-saturation parameter (I_k) was redundant and potentially misleading. Although U. rotundata exhibits a great amplitude of photoacclimation, it apparently has little capacity for temperature acclimation compared to the kelp, Laminaria saccharina, for which published data indicate similar photosynthetic rates over a broad range of growth temperatures. Diurnal variation of P_m and R_d at a growth PFD of ～ 1700 ± 200 μmol photons · m^?2· s^?1 was similar to the pattern observed previously in outdoor culture, suggesting endogenous control of these parameters. Quantum yield and the ratio of variable to maximum chlorophyll fluorescence (F_v/F_m), which were depressed in midday sunlight exceeding ～ 1500 μmol photons · m^?2· s^?1, were relatively invariant through the day in indoor culture, indicating that these parameters are controlled primarily by instantaneous PFD. Growth and fluorescence data are also presented for some other macroalgae for comparative purposes.     

EFFECTS OF TEMPERATURE,NITROGEN SUPPLY,AND TISSUE NITROGEN ON AMMONIUM UPTAKE RATES OF THE CHLOROPHYTE SEAWEEDS ULVA CURVATA AND CODIUM DECORTICATUM1

Nitrogen uptake rates of Ulva curvata (Kütz.) de Toni (Ulvales) and Codium decorticatum (Woodw.) Howe (Caulerpales) grown under several N addition regimes were determined by perturbation and continuous mode techniques, and as N demand, by the product of growth rate and tissue N. Uptake rates are reported as the slope of rate vs. concentration curves in each case. N uptake rates of U. curvata were inversely correlated with tissue N and affected only slightly by temperature. There was no correlation of N uptake rate with tissue N in C. decorticatum. N uptake rates of C. decorticatum were affected by temperature but to a lesser degree than were growth rates. Neither N addition per se nor light affected N uptake capacity of either species. The proximal mechanism for seaweeds accumulation of N at low light and temperatures may be that N uptake is less limited by light and temperature than is growth. This in turn may partially compensate for the effects of reduced light and temperature on growth by increasing pigment and enzyme levels. Perturbation uptake rates were higher than continuous mode or N demand rates in Ulva but not in Codium. N uptake rates of Ulva were higher than those of Codium, but N storage capacities were lower. These two observations suggest that Ulva experiences a fundamentally more variable N supply than does Codium. This is consistent with the clarification of Ulva as an ephemeral form and of Codium as persistent. A seaweed's functional form therefore appears to influence the spectrum of resource variability available to it as well as its ability to persist in the environment.     

THE RED ALGAL EPIPHYTES MICROCLADIA COULTERI AND M. CALIFORNICA (RHODOPHYCEAE,CERAMIACEAE). II. BASIPHYTE SPECIFICITY1

The phenomenon of basiphyte specificity in the settlement and growth of the red algal epiphytes Microcladia californica Farl. and M. coulteri Harv. was examined by studying the interface with their respective basiphytes and by cross-inoculation experiments. Microcladia californica attaches only to the surface of its single basiphyte Egregia menziesii (Aresch.) Turn. whereas M. coulteri penetrates the tissue of a wide range of basiphytes. The pattern of primary rhizoid development in both epiphytes determines the mode of attachment and may influence the range of basiphytes possible for each epiphyte. Cross-inoculation experiments show that Microcladia californica is not able to colonize the basiphytes of M. coulteri, Iridaea and Prionitis, or Ulva. The mechanisms by which these algae restrict the growth of epiphytes include short life-span, “cuticle peeling” and chemical defense. Microcladia coulteri, which naturally colonizes Iridaea and Prionitis, has evolved mechanisms to counteract the antifouling effects of those basiphytes. The question of why Egregia is the exclusive substratum for M. californica remains undetermined. However, Egregia may provide the appropriate ecological conditions and a surface topography conductive to M. californica spore settlement and growth.     

PHOTOSYNTHETIC PROPERTIES OF PROTOPLASTS,AS COMPARED WITH THALLI,OF ULVA FASCIATA (CHLOROPHYTA)1

Protoplasts were prepared from Ulva fasciata Delile, and their photosynthetic performance was measured and compared with that of thalli discs. These protoplasts maintained maximal rates of photosynthesis as high as those of thalli (up to 300 μmol O₂·mg chlorophyll^?1·h^?1) for several hours after preparation and were therefore considered suitable for kinetic studies of inorganic carbon utilization. The photosynthetic K_1/2(inorganic carbon) at pH 6.1 was 3.8 μM and increased to 67, 158, and 1410 μM at the pH values 7.0, 7.9, and 8.9, respectively. Compared with these protoplasts, thalli had a much lower affinity for CO₂ but approximately the same affinity for HCO₃^?. Comparisons between rates of photosynthesis and the spontaneous dehydration of HCO₃^? (at 50 μM inorganic carbon) revealed that photosynthesis of both protoplasts (which lacked apparent activity of extracellular/surface-bound carbonic anhydrase) and thalli (which were only 25% inhibited by the external carbonic anhydrase inhibitor acetazolamide) could not be supported by CO₂ formation in the medium at the higher pH values, indicating HCO₃^? uptake. Since both protoplasts and thalli were sensitive to 4,4′-diisothiocyanostilbene-2,2′-disulfonate, we suggest that HCO₃^? transport was facilitated by the membrane-located anion exchange protein recently reported to function in certain Ulva thalli. These findings suggest that the presence of a cell wall may constitute a diffusion barrier for CO₂, but not for HCO₃^?, utilization under natural seawater conditions.     

Novel distribution pattern between coexisting sexual and obligate asexual variants of the true estuarine macroalga Ulva prolifera

Where sexual and asexual forms coexist within a species, the asexuals are often found to be prevalent in marginal habitats. This asexual distribution pattern has received evolutionary attention linked to the paradox of sex. In many marine species, there is a distributional trend of asexual modes being more common in lower salinity waters regarded as the ecogeographic marginal, being explained by negative effects of low salinities on sexual reproductive success. However, the distribution pattern of estuarine species recently adapted to low salinity waters has remained unknown. The brackish macroalga Ulva prolifera being a major benthic component of estuarine ecosystems includes a sexual variant and obligate asexual variants by means of motile spores. We examined the sexual–asexual distribution pattern of this alga along a salinity gradient in river estuaries. Surprisingly, opposite to the distributional trend of marine organisms, the results clearly showed the persistent predominance of sexuals in the lower salinity reaches than the asexuals. In addition, a frequent alternating of dioecious gametophytes and sporophytes in the sexual population was observed, suggesting the sexual reproductive process would be robustly performed in the low salinity waters. Considering U. prolifera had evolved from the ancestral marine species to become a true estuarine species of which the core habitat is the low salinity reaches, in a broad sense its sexual–asexual distribution pattern would be involved in asexual marginal occupations of the species range previously reported in other organisms. Based on the frozen niche variation model, we can give a concise explanation for the evolutionary process of this pattern; multiple asexuals with frozen genotypic variation had arisen from sexual ancestors undergoing low salinity adaptation and share the estuarine habitat with the sexuals at present.     

Examination of prezygotic and postzygotic isolating barriers in tropical Ulva (Ulvophyceae,Chlorophyta): evidence for ongoing speciation

Phylogenetic clades based on DNA sequences such as the chloroplast rbcL gene and the nuclear ITS region are frequently used to delimit algal species. However, these molecular markers cannot accurately delimit boundaries among some Ulva species. Although Ulva reticulata and Ulva ohnoi occasionally bloom in tropical to warm‐temperate regions and are clearly distinguishable by their reticulate or plain blade morphology, they have few or no sequence divergences in these molecular markers and form a monophyletic clade. In this study, to clarify the speciation and species delimitation in the U. reticulata‐ohnoi complex clade, reproductive relationships among several sexual strains from the Philippines and Japan including offspring that originated from the type specimen of U. ohnoi were examined by culturing and hybridization in addition to the ITS‐based analysis. As a result, both prezygotic and postzygotic reproductive isolation were revealed to occur between genetically perforated U. reticulata and imperforate U. ohnoi. They were also separated on the basis of sequence analysis of the ITS region. That strongly supports that the two taxa are independent biological species. Although no prezygotic barrier among the Philippine and Japanese strains of U. reticulata was observed, unexpectedly zoospores produced by hybrid sporophytes in some of their combinations mostly failed to develop, indicating partial formation of a postzygotic barrier despite a 0.2% divergence in the ITS sequence. These findings suggest speciation is still ongoing in U. reticulata.     

24-表油菜素内酯和盐度对浒苔生长和生理活性的影响

为探讨低温条件下外源植物激素和盐度变化对浒苔光合生理的影响,本文选取浒苔为供试材料,研究24-表油菜素内酯和盐度对其生长、光合色素、叶绿素荧光参数、抗氧化活性和可溶性蛋白含量的影响.结果表明: 与正常盐度(近岸海水盐度为25)相比,浒苔的生长在盐度10处理下显著增加,增幅为45.9%,但在盐度5处理下显著降低.盐度5处理的浒苔比其他盐度处理具有较高的Chl a和可溶性蛋白.24-表油菜素内酯(0.2 mg·L^-1)的加入显著抑制了浒苔的生长,尤其在盐度25处理下,浒苔呈现负增长的趋势,并大量释放孢子(配子),有效光化学效率、超氧化物歧化酶酶活性和可溶性糖含量明显降低,但可溶性蛋白含量显著增加.即在温度15 ℃条件下,可通过适当降低盐度促进浒苔的生长;在盐度25条件下,可通过添加24-表油菜素内酯促进孢子(配子)释放,为浒苔的大规模养殖提供原材料.     

Flagellar scales in Oxyrrhis marina Dujardin

Oxyrrhis marina Dujardin possesses imbricated scales of a tight spiral form on both flagella. These and the associated flagellar hairs are described in detail.