PATTERNS OF RELATIVE GROWTH IN NEREOCYSTIS LUETKEANA (PHAEOPHYTA)1

Plants of Nereocystis luetkeana (Mert.) Post. et Rupr. of different sizes were held on a raft at the surface of the sea, off the Friday Harbor Laboratories, San Juan Island, Washington for 5-day periods for obseruation of detailed relative growth of different parts. Each stipe was marked at intenlals with injected Indian ink and each blade was punched with a series of holes. Measurements of diameter, length, width and thickness were made before and after the 5-day periods. Blades showed a very similar pattern of relative growth rate (R) ovler an 18-fold range of sizes. The maximum local R in length was about 0.2 day⁻¹ and occurred at 6.5% of the distance from the bulb to the tip, declining to 0.01 at half way. Half the linear growth occurred in the proximal one tenth of the blade and 95% within the proximal half: The relative growth rate of the whole blade declined only slightly with increased size and lay between 0.03 and 0.06 day⁻¹ (approx. 3-6% day⁻¹). The linear growth rate therefore increased with blade size, the maximum observed being 14 cm day⁻¹. The maximum relative growth rate in blade width was slower, and sited more distally than that in length. Unless fertile tissue was involved all blade tissue, except that closely adjoining the bulb, became thinner duringgrowth. R in volume reached 0.3 day⁻¹. Presumably because the plants were held near the sea surface stipes grew slowly, with a maximum linear rate of 9 mm day⁻¹. The maximum R in length decreased with stipe length. Bulb R in volume also decreased as size increased, from a maximum of 0.3 day⁻¹.     

GROWTH OF BLADES OF NEREOCYSTIS LUETKEANA (PHAEOPHYTA) IN DARKNESS 1

Short term measurements were made of the relative growth rate of the fast growing portions of blades of Ncreocystis luetkeana (Merl.) Post. et Rupr. during exposure to natural daylight and during prolonged darkness. Growth was only slightly, but significantly, faster during the 12 h of daylight than during the 12 h that included 8 h of darkness. Clearly considerable growth occurred at night. In blades amputated 1 cm beyond the zone measured, growth was slower during both night and day. In continuous darkness growth continued for up to 12 days. It was not influenced by amputation of distal blade tissue but it was increased by severance from the bulb and stipe. A mean volume increase of 50% of the blade tissue was recorded. While the organic content decreased, the drop was half that required to support the increase in volume. There was some evidence against translocation. It is possible that cellular biochemical rearrangements allow a light-independent increase in organic material.     

SEASONAL VARIATIONS IN TOTAL NITROGEN,CARBON AND AMINO ACIDS IN MACROCYSTIS INTEGRIFOLIA AND NEREOCYSTIS LUETKEANA (PHAEOPHYTA)1

Seasonal variations in tissue nitrogen, carbon, amino acids and ammonium were determined for the brown algae Macrocystis integrifolia Bory and Nereocystis luetkeana (Mertens) Pastels and Ruprecht, For M. integrifolia, the proportions of tissue nitrogen and carbon in blades, bulbs and stipes were also determined. The composition of the two algae in terms of the above constituents was similar. In addition, ammonium, nitrogen and protein-bound amino acids showed distinct seasonal trends with high values during the winter and low levels during the summer. The range for nitrogen was 0.8–3.0% and for proteins 7.6–11.7% of dry weight. In contrast, carbon content and C/N ratio showed the reverse trend with higher values during the summer and lower values during the winter. The range for carbon was 19–31% of dry weight, and the C/N ratio showed a range of 9–37. The free amino acids did not show any specific seasonably. Tissue nitrogen and carbon showed higher values in the blades than in the bulbs and stipes.     

HYBRIDIZATION OF MACROCYSTIS (PHAEOPHYTA) WITH OTHER FLOAT-BEARING KELPS1,2

Sporophytes were produced in 22 out of 23 attempts to cross Macrocystia angustifolia Bory with Pelagophycus porra (Lemon) Setchell. Representative samples of these plants were raised in a surge-tank under greenhouse conditions to several meters in length. The plants resulting from this intergeneric cross were intermediate in morphology between the parental genera. The intermediate plants were similar to those found fry one of the authors (MN) in 1957, and by others since then, in California kelp beds where Macrocystis and Pelagophycus co-occur, A cross between M. angustifolia and Nereocystis luetkeana (Mertens) P. & R., also produced normal sporophytes. The taxonomic, evolutionary, morphogenetic and genetic implications of these hybridizations are discussed.     

CIRCADIAN GROWTH RHYTHM IN JUVENILE SPOROPHYTES OF LAMINARIALES (PHAEOPHYTA)1

A circadian rhythm in growth was detected by computer-aided image analysis in 3–4-cm-long, juvenile sporophytes of the kelp species Pterygophora California Rupr. and in seven Laminaria spp. In P. californica, the free-running rhythm occurred in continuous white fluorescent light, had a period of 26 h at 10°or 15°C, and persisted for at least 2 weeks in white or blue light. The rhythm became insignificant in continuous green or red light after 3 cycles. Synchronization by white light-dark regimes, e.g. by 16 h light per day, resulted in an entrained period of 24 h and in a shift of the circadian growth minimum into the middle of the light phase. A morning growth peak represented the decreasing portion of the circadian growth curve, and an evening peak the increasing portion. The circadian growth peak was not visible during the dark phase, because growth rate decreased immediately after the onset of darkness. At night, some growth still occurred at 16 or 12 h light per day, whereas growth stopped completely at 8 h light per day, as in continuous darkness. During 11 days of darkness, the thallus area became reduced by 3.5%, but growth rate recovered in subsequent light–dark cycles, and the circadian growth rhythm reappeared in subsequent continuous light.     

ETIOLOGY AND DEVELOPMENT OF HYPERPLASIA INDUCED BY STREBLONEMA SP. (PHAEOPHYTA) ON MEMBERS OF THE LAMINARIALES (PHAEOPHYTA)1

Filaments of Streblonema sp. isolated from hyperplasia (galls) on Nereocystis luetkeana (Mert.) Post. et Rupr. induced similar gall growths when inoculated on young sporophytes of Nereocystis luetkeana, Macrocystis integrifolia Bory, and Laminaria japonica Aresch. Filaments were irregularly branched and uniseriate with cells 5–8 μm in diameter and 10–30 μm long. Growth under varied cultured conditions produced unilocular sporangia. Zoospores released from the sporangia germinated into identical filaments which also formed unilocular sporangia. Gall tissue originated from the innermost cells of the epidermal meristematic zone. When infected these cells divided in an irregular and unorganized manner. The resulting structure was a pronounced departure from normal morphology.     

PHOSPHORUS AND THE GROWTH OF JUVENILE MACROCYSTIS PYRIFERA (PHAEOPHYTA) SPOROPHYTES1

The effect of phosphate (P_i) supply on growth rate and tissue phosphorus content of juvenile Macrocystis pyrifera (L.) C. Ag. sporophytes was examined. Sporophytes were batch cultured in aquaria with flowing recirculated seawater enriched by 30 μM nitrate. Each aquarium was supplemented with a different seawater P_i concentration, 0, 0.3, 1, 2, 3, and 6 μM. Sporophyte mean specific growth rates declined with time in all cultures presumably due to the normal developmental decrease in the proportion of meristematic tissue of each plant. Growth rate declines were more pronounced in cultures that were nutrient limited. Sporophyte growth was P-limited after two-week exposure to P_i less than 1 μM, corresponding to a tissue P concentration of less than 0.20% dry weight. Plants cultured at 6 μM P_i contained tissue P levels of 0.53% dry weight after three weeks. Luxury consumption and storage of P occurred.     

COLONIZATION AND GROWTH OF ASCOPHYLLUM NODOSUM (PHAEOPHYTA) IN MAINE1

Colonization and growth of Ascophyllum nodosum (L.) Le Jol. were investigated in two Maine estuaries from 1972 to 1978. Following denudation of intertidal rock, substrata were initially colonized by Fucus vesiculosus L.; eventually, Ascophyllum supplanted Fucus, and became dominant in terms of percentage cover. Ascophyllum settled first and most densely in the low intertidal zone, but its fastest growth occurred in the mid-intertidal zone. Some, but not all, Ascophyllum germlings produced a vesicle within one year of colonization. The mean annual growth of A. nodosum was variable among sites, zones and years.     

PHYTOCHROME-MEDIATED STIPE ELONGATION IN THE KELP NEREOCYSTIS (PHAEOPHYCEAE)1

Preliminary evidence is presented which indicates that far-red irradiation promotes and near-red inhibits rapid stipe elongation in Nereocystis luetkeana (Mertens) Postels & Ruprecht. Laboratory results showed afar-red reversal of red inhibition, but a red/far-red reversal was not evident in field experiments. The effects of changing ratios of red and far-red irradiation brought about by selective water absorption are discussed in relation to stipe elongation of Nereocystis.     

SEASONAL PATTERNS OF MORPHOLOGICAL VARIABILITY IN SARGASSUM POLYCERATIUM (PHAEOPHYTA)1

Temporal variability in certain morphological and taxonomically important features was quantified for Sargassum polyceratium Mont. from a population in the Content Keys, Florida (U.S.A.). Patterns of blade development, senescence, and loss caused pronounced seasonal changes in blade length-width ratios. Blade length and width were maximal early in the growing season (August-November) and decreased as the annual stems matured. Early in the growing season, plants had broader blades with randomly distributed cryptostomata. Late in the growing season (February-April), plants had more linear blades with cryptostomata approximately arranged in two rows, one on each side of the midrib. The length-width ratio of blades increased acropetally along the stems and were directly correlated to the size of the cryptostomatal opening and inversely correlated with the number of cryptostomata. The branching pattern of the annual stems ranged from short spur branches to well-developed, lateral axillary branches. The frequency of bifurcated blades increased significantly late in the growing season. Vesicle shape and size and pedicel length were temporally stable. Alated pedicels and mucronate vesicles occurred in low frequencies. The variability of the morphological features used to delineate species within the genus Sargassum on the tropical eastern coasts of the Americas is poorly understood.     

腕足类的相关生长与统计鉴别

采用生物统计法中常用的双变量分析,研究腕足类种一级分类单位。选取壳长和壳厚作为一对相关变量,通过对生长线斜率和位置的假设检验,判断同属异种之间是否存在实质性差别。建立一个新种时,必须确认其相关生长线（ｌｉｎｅ ｌｆ ｒｅｌａｔｉｖｅ ｇｒｏｗｔｈ）具有正常的形式;标本残破不易正确划分时,可计算双变量的相关系数,预测样品是否划归不同的种。     

ADAPTATION OF PHOTOSYNTHESIS IN LAMINARIA SACCHARINA (PHAEOPHYTA) TO CHANGES IN GROWTH TEMPERATURE1

The effect of growth temperature on photosynthetic metabolism was studied in the kelp Laminaria saccharina (L.) Lamour. Photosynthesis was subject to phenotypic adaptation, with almost constant photosynthetic rates being achieved at growth temperatures between 0 and 20° C. This response involved: (1) an inverse relationship between growth temperature and photosynthetic capacity, (2) a reduction in the Q₁₀ value for photosynthesis of L. saccharina grown at 0 and 5° C compared with 10, 15 and 20° C grown sporophytes, and (3) an acquired tolerance of photosynthesis to temperatures between 15–25° C (which inhibited photosynthesis in 0 and 5° C grown L. saccharina) in sporophytes grown at 10, 15 and 20° C. The physiological basis of these adaptations is discussed in terms of observed changes in activities and kinetics of the Calvin cycle enzyme ribulose-1, 5-bisphosphate carboxylase (oxygenase) and efficiency of light harvesting-electron transport systems.     

PATTERNS OF GROWTH AND CELL COMPOSITION OF FRESHWATER ALGAE IN LIGHT-LIMITED CONTINUOUS CULTURES1

Selenastrum minutum (Näg.) Collins and three species of Scenedesmus were grown in light-limited continuous cultures. Comparative growth patterns were explained in terms of differences in the rates of light absorption and utilization. For all species, increasing light intensity or lowering temperature caused chl a/C to decrease and RNA/C to increase. Similar shifts in pigment composition have been widely reported. However, the RNA:light interaction has only been reported infrequently, and the RNA:temperature interaction is believed to be the first reported observation of its kind in algae. Distinct differences in RNA/C among the test algae may be size-dependent. Similar environmentally induced shifts in chl a/C and RNA/C among all test algae point to the existence of a common control mechanism which maximizes the specific growth rate through adjustments to cell composition.     

CIRCANNUAL GROWTH RHYTHM AND PHOTOPERIODIC SORUS INDUCTION IN THE KELP LAMINARIA SETCHELLII (PHAEOPHYTA)1

Growth and reproduction of laboratory-grown sporophytes of Laminaria setchellii Silva were investigated in a tank system with controlled conditions of daylength, temperature, and nutrients (N and P). A circannual growth rhythm of the frond was detected under constant laboratory conditions. In continuous long-day and night-break conditions the period τ of the free-running rhythm varied between 11.3 and 17.3 months; in short-day conditions the frond grew indefinitely. The growth rhythm of individual plants could be synchronized by a simulated annual cycle of day-length with a period of T = 12 months. The four seasons of the year were simultaneously simulated by phase shifting the annual cycle of daylength by 3, 6, or 9 months in three out of four tanks. The annual growth cycle followed these phase shifts, and initiation of the new blade always started just after the winter daylength minimum. The formation of sori was induced by a genuine photoperiodic short-day reaction in 1- to 2-year-old plants. Sori became, visible 9–14 weeks after transfer of individual plants from long-day to short-day conditions, whereas plants cultured in continuous long-day or night-break conditions remained sterile. Sporophytes with or without blades were able to continue growth or produce new blades in continuous darkness.     

EFFECTS OF NITRATE CONCENTRATION ON THE GROWTH AND PHYSIOLOGY OF LAMINARIA SACCHARINA (PHAEOPHYTA) IN CULTURE1,2

Laminaria saccharina Lamour. sporophytes were grown in enriched and synthetic media through a range of nitrate concentrations, There was an approximately linear relationship between growth and nutrient concentration up to 10 μ substrate concentration. The half-saturation constant (K_{2) was ca. 1.4 μ NO3-. The internal levels of NO3- increased at substrate concentrations above 10 μM b>3- and reached levels several thousand times higher than the surrounding medium. Thus there is evidence for luxury consumption of NOsb>3}^-. The chlorophyll content and photosynthetic capacities of plants also increased with increasing external NO₃^- The ecological implications of this work are considered.     

DAYLENGTH AND LIGHT RESPONSES IN GROWTH AND FERTILITY OF GLOSSOPHORA KUNTHII (PHAEOPHYTA,DICTYOTALES) FROM PACIFIC SOUTH AMERICA1

Excised ligulae of Glossophora kunthii (C. Ag.) J. Ag. were cultured in photoperiods of 4–24 h and photon fluence rates of 10–75 μmol.m^?2.s^?1. Daylength interacted with irradiance on the growth of the ligulae. Maximal growth of primary ligulae occurred in long-day regimens with high irradiances suggesting an effect of irradiance on photosynthesis and growth. In contrast, growth of secondary ligulae was greatest in short-day regimes. Differences were significant at the highest irradiance tested. Differentiation of tetrasporangia on the ligulae is a short-day photoperiodic response. Daylengths of 8.5 h or less induced a sharp increase in numbers of fertile ligulae and tetrasporangia attaining maturity. Interruptions of the dark period decreased the development of tetrasporangia; the number of interruptions had a cumulative inhibitory effect. Differentiation of reproductive structures was influenced by interactions of photoperiod and irradiance. Maximum numbers of tetrasporangia were formed at short-day regimes and low irradiances; differentiation was completely inhibited at long-day conditions and high irradiance.     

SEASONAL MODULE DYNAMICS IN SARGASSUM SUBREPANDUM (FUCALES,PHAEOPHYTA)1

Module dynamics of the fucoid alga SARGASSUM SUBREPANDUM (Forssk.) C. Agardh was studied in the southern Red Sea. Seasonal variation in thallus density and size was determined, and the initiation, growth, reproduction, and shedding of modules (primary laterals) were ascertained, using a tagging approach. Possible effects of different size‐related parameters on module initiation, growth, reproduction, and shedding were analyzed in the context of contradicting results for other macroalgae, in comparison with terrestrial plants. Thallus density varied little; most of the seasonal variation occurred at the modular level. A restricted period of new module formation early in the cooler season was followed by fast growth and reproduction. Massive shedding of modules occurred toward the end of the cooler season leading to strongly reduced biomass in summer. There was some evidence that high module numbers inhibited new module formation and enhanced the maximum module elongation rate (fastest‐growing module per thallus). On the other hand, elongation rates generally decreased, and apical tissue losses increased with increasing module length. This response was observed over a wide size range, suggesting grazing losses. There was no evidence of suppressed growth in small modules due to intraspecific competition. Elongation rates remained unaffected by reproductive status, indicating that there was no direct trade‐off between growth and reproduction. Module survivorship was independent of module number and size, but fertile modules were more persistent than vegetative ones. We conclude that module dynamics are determined by seasonal changes in the environment, size‐dependent processes, and interactions among the modules.     

VARIATION IN NITROGEN PHYSIOLOGY AND GROWTH AMONG GEOGRAPHICALLY ISOLATED POPULATIONS OF THE GIANT KELP,MACROCYSTIS PYRIFERA (PHAEOPHYTA)1

Three geographically isolated populations of the giant kelp, Macrocystis pyrifera (L.) C. Ag., were examined for responses to nitrate availability in batch culture experiments using juvenile sporophytes reared from spores in the laboratory. Although maximum rates of nitrate-saturated growth were similar among groups, there were significant quantitative differences in the response to nitrate limitation that can be related to natural patterns of nutrient availability at these sites. Plants from Santa Catalina Island (most oligotrophic) achieved maximum growth rates at ambient nitrate concentrations that were lower than those for plants from Monterey Bay, California (most eutrophic), or Refugio State Beach (near Santa Barbara, California). Tissue nitrogen and amino acid concentrations were highest in plants cultured from Santa Catalina Island populations at all external nitrate concentrations, suggesting that differences in nitrate requirements for growth may reflect the efficiency of nitrate uptake and assimilation at subsaturating nitrate concentrations. Given the different physical environments from which these plants came, the data suggest that geographically isolated populations of M. pyrifera have undergone genetic divergence that can be explained by ecotypic adaptation to unique habitat conditions at these sites.     

GROWTH AND PRODUCTION OF LAMINARIA LONGICRURIS (PHAEOPHYTA) POPULATIONS EXPOSED TO DIFFERENT INTENSITIES OF WATER MOVEMENT1

Blade elongation was compared in two populations of Laminaria longicruris de la Pylaie in Shag Bay, Nova Scotia, Canada; one population was exposed to and the other sheltered from high intensity water movement. The maximum and minimum elongation rates were similar for the two populations, but the sheltered coast plants grew more rapidly during 8 mo of the year. Measurements of internal inorganic nitrogen (NO₃^?+ NO₂^?) reserves and dry organic weight indicated that low concentrations of dissolved nutrients in summer and low levels of illumination in fall and winter were more limiting to growth at the exposed site than the sheltered site. Transplant experiments provided evidence that morphological differences between the exposed and sheltered coast plants accounted at least partially for their different responses to varying nutrient conditions and light levels. It is concluded that lower productivity of the exposed coast population is the result of adaptation to high intensity water movement.     

LIGHT DEPENDENCE OF GROWTH AND PHOTOSYNTHESIS IN PHAEODACTYLUM TRICORNUTUM (BACILLARIOPHYCEAE)1

Phaeodactylum tricornutum Bohlin was maintained in exponential growth over a range of photon flux densities (PFD) from 7 to 230 μmol·m^?2s^?1. The chlorophyll a-specific light absorption coefficient, maximum quantum yield of photosynthesis, and C:N atom ratio were all independent of the PFD to which cells were acclimated. Carbon- and cell-specific, light-satuated, gross photosynthesis rates and dark respiration rates were largely independent of acclimation PFD. Decreases in the chlorophyll a-specific, gross photosynthesis rate and the carbon: chlorophyll ratio and increases of cell- or carbon-specific absorption coefficients were associated with an increase in cell chlorophyll a in cultures acclimated to low PFDs. The compensation PFD for growth was calculated to be 0.5 μmol·m^?2s^?1. The maintenance metabolic rate (2 × 10^?7s^?1), calculated on the basis of the compensation PFD, is an order of magnitude lower than the measured dark respiration rate(2.7 × 10^?6mol O₂·mol C^?1s^?1). Maintenance of high carbon-specific, light-saturated photosynthesis rates in cells acclimated to low PFDs may allow effective use of short exposures to high PFDs in a temporally variable light environment.