PHYSIOLOGICAL INVESTIGATIONS ON ALARIA ESCULENTA (LAMINARIALES,PHAEOPHYCEAE). II. ROLE OF TRANSLOCATION IN BLADE GROWTH1,2

Laboratory studies on blade growth in Alaria esculenta (L.) Grev. showed 3 periods of rapid blade elongation during the year: October–November, February–April and late June. The first two periods are characteristic of many Laminariales; the unique June peak may reflect local nutrient conditions. While the distal blade functions as a source, supplying organic matter to the blade meristem, the stipe can be a source during periods of rapid growth or a sink during late summer when blade growth is slow. Maximum enhancement of elongation rate of blade meristems was observed in 40–50 cm blades; longer blades showed no further increase in growth rate. This blade length-growth promotion relationship may be independent of seasonal variations in meristematic activity. ¹⁴C tracer experiments suggested that separate growth promotion effects by distal blade, sporophylls and stipe were not additive in the intact thallus. The preferential source of assimilate for blade meristem growth was the distal blade. Secondary sources: sporophylls, which were activated following excision of the primary source; and stipe, which began to translocate assimilate when both sources were removed. The role of secondary sources in nature is discussed. Profiles of radioactivity in alcohol-soluble organic matter in blades are evaluated in relation to tracer profiles in higher plants and mechanisms of translocation.     

Relation between size and age of holdfasts of Ecklonia stolonifera Okamura (Laminariales,Phaeophyta) in northern Honshu,Japan

Ecklonia stolonifera is distributed along the coast facing the Sea of Japan. The size of various parts of the shoot (blade length and width and stipe length and diameter) and the age were determined at Ooma, Aomori Prefecture. The smaller the holdfast, the higher the percentage of one-year-old shoots. Holdfasts 10 cm in diameter seemed to be three years old, whereas holdfasts 40 cm in diameter seemed to be five or more years old. Zoosporangial sori were observed on blades three or more years old. Ecklonia stolonifera holdfast diameter expands only vegetatively by stoloniferous rhizoids. Zoospores, formed on shoots three or more years old, serve for the formation of new populations.     

DEVELOPMENTAL STUDIES IN PORPHYRA. I. BLADE DIFFERENTIATION IN PORPHYRA PERFORATA AS EXPRESSED BY MORPHOLOGY,ENZYMATIC DIGESTION,AND PROTOPLAST REGENERATION1

Four areas containing different cell morphologies were mapped on Porphyra blades and five different cell types (i.e. tapered with long extensions, large and vacuolated, vegetative and dividing, and reproductive: males and females) were identified in them. Tissues from these areas were dissociated, and protoplasts and single cells were isolated from the dissociated tissue of each distinct region. Regeneration rates of the isolated cells and protoplasts (isolates) varied depending on their morphological type. Regeneration rates were lowest in cultured isolates from the area just above the holdfast (ca. 1 %) and increased gradually to over 80% in isolates from areas of vegetative and reproductive regions away from the holdfast. Four distinct morphological patterns were observed among the regenerating plants. Cells isolated from vegetative areas developed into leafy plants while in liquid culture, and into calli when grown on solid medium. Isolates from reproductive areas developed into either a long thin or short thick filamentous plant. Those from ripe patches of carposporangia developed into thin conchocelis filaments, while isolates from non-differentiated cells bordering the ripe reproductive patches developed into thick filaments resembling the morphology of conchosporangial branches. The blade of Porphyra appears simple as it consists of a single cell layer; however, it is complex both morphologically and physiologically.     

MORPHOLOGY AND TAXONOMY OF HIMANTOTHALLUS (INCLUDING PHAEOGLOSSUM AND PHYLLOGIGAS), AN ANTARCTIC MEMBER OF THE DESMARESTIALES (PHAEOPHYCEAE)1

The sporophyte of Himantothallus develops according to a closed pattern in which the number and position of the blades is determined by the location of trichothallic meristems in a filamentous germling. Expansion of the miniature juvenile to the massive adult thallus is accomplished by diffuse secondary growth and involves a change from filamentous rhizoids to a hapteroid holdfast, flattening of the stipe, and enormous increases in length, breadth, and thickness of both stipe and blade. The axis usually bears 1–8 lateral blades, often paired, and terminates in a flattened stub. Phaeoglossum is interpreted as a growth form of Himantothallus in which a terminal blade develops to the exclusion of lateral blades, the latter being represented by a single spine. Phyllogigas clearly falls within the morphological spectrum of Himantothallus, the lack of twisting being related to physical factors in the environment. Sporangia, interspersed with an equal or somewhat larger number of two-celled paraphyses, are borne in slightly elevated sori scattered over both surfaces of the blade. Zoospore germination was not observed, nor were gametophytes, either in culture or in the field. Haptera apparently originate from the meristoderm in the lower part of the maturing stipe and lack a filamentous medulla. The mature stipe and the mature blade are anatomically similar, being composed of a superficial meristoderm, a cortex of parenchyma-like cells, and a filamentous medulla. The meristoderm is usually a single layer of plastid-containing cells that divide anticlinally to accommodate (or effect) expansion and periclinally to produce cortical tissue inward. Cortical cells are in radial files and increase in diameter towards the interior. They usually are densely packed with physodes. The medulla is uniquely distinguished by the presence of sheathed trumpet hyphae. Cells of the trumpet hyphae have perforate end walls with callose deposits and probably function in conduction as do the sieve filaments in Laminariales. Sheathing cells are filled with plastids. Sheathing filaments form connections among themselves and with nearby unsheathed filaments. The sheathed trumpet hyphae and their matrix of unsheathed filaments form a plexus, which in the mature blade is flattened and may be stripped intact from the other tissues. Development of the embryonic sporophyte is very similar to that in Desmarestia, as is the anatomy of the adult thallus and the sporangia. From these considerations, Himantothallus is assigned to the Desmarestiaceae (Desmarestiales).     

CARBON ALLOCATION IN MACROCYSTIS PYRIFERA (PHAEOPHYTA): INTRINSIC VARIABILITY IN PHOTOSYNTHESIS AND RESPIRATION1

Measurements of net photosynthesis (PS, O₂ evolution), dark respiration (R, O₂ consumption), and light and dark carbon fixation (¹⁴C) were conducted on whole blades, isolated blade discs, sporophylls, apical scimitars and representative portions of stipe and holdfast of the giant kelp Macrocystis pyrifera L.C. Ag. On a dry weight basis, highest net PS rates were observed in apical scimitar segments and whole blades (3.81 and 3.07 mgC · g dry wt^?1· h^?1, respectively), followed by sporophylls (1.42 mgC·g dry wt^?1· h^?1) and stipe segments (0.15 mgC·g dry wt^?1· h^?1). No PS capacity was observed in holdfast material. Respiration rates showed similar ranking ranging from 1.22 mgC·g dry wt^?1·h^?1 for apical scimitar to 0.18–0.22 mgC·g dry wt^?1· h^?1 for holdfast material. Considerable within blade variability in both PS and R was also found. Steepest PS and R gradients on both an areal and weight basis were found within immature blades followed by senescent and mature blade material. Highest net PS rates were associated with the blade tips ranging from 3.08 (mature blades) to 10.3 mgC·dry wt^?1·h^?1 (immature blades). Highest rates of R generally occurred towards the basal portions of blades and ranged from 1.03–1.80 mgC·g dry wt^?1·h^?1 for immature blades. The variability within and between blades was high, with coefficients of variation approaching 50%. The observed patterns can be related to the decreasing proportionment of photosynthetic tissue and increasing proportionment of structural tissue as occurs from the blade tip to the blade base. Rates of light carbon fixation (LCF) revealed longitudinal profiles similar to oxygen measurements for the different blade types, with the absolute rates being slightly lower. Patterns of dark carbon fixation (DCF) were less easily interpreted. Highest rates of DCF (0.04–0.06 mgC·g dry wt^?1·h^?1) occurred at the basal portions of immature and senescent blades. Longitudinal profiles of total chlorophyll (a + c) on both an areal and weight basis were very similar to the profiles of PS. Normalized to chlorophyll a, PS displayed an unusual longitudinal profile in immature tissue; however, such profiles for mature and senescent tissues were similar to those for PS on an areal basis. It was demonstrated that it is difficult, if not impossible, to select single tissue discs that are representative of whole blades. The metabolic longitudinal profiles reveal a characteristic developmental pattern; the previous working definitions of immature, mature, and senescent blades, based on morphology and frond position thus have a physiological basis.     

The life history of Petrocelis franciscana

Tetraspores from Petrocelis franciscana Setch. et Gardn. collected at Rockaway Beach, San Mateo County, California, 11 January 1971 were isolated into unialgal culture with Provasoli's enriched seawater medium at 10° and 15° C, 16:8 daily photoregime, 2000–2500 lx cool white fluorescent lighting. The germlings developed as crustose discs with marginal meristems until about 300–500 µm in diameter when erect multiaxial blades were established. The erect plants grew to reproductive maturity in 15° C but not in 10° C. By the fifth month of growth the erect axes flattened and branched dichotomously. At the end of 7 months the plants were reproductively mature. These cultured plants were morphologically similar to Gigartina agardhii Setch. et Gardn. (subgenus Mastocarpus) which occurs abundantly at Rockaway Beach. Male plants were non-papillate with spermatangia forming on the entire blade surface except at the growing tip and near the stipe. The female plants developed papillae which bore many procarps. Many of these procarps were penetrated and seemingly destroyed by intrusive filaments from adjacent vegetative branches but apparently a number were functional since a number of cystocarps developed on plants in both stationary and shake culture after 5 months. Some viable spores were released and germinated.     

CONTRASTING PATTERNS OF ALLOMETRY AND REALIZED PLASTICITY IN THE SISTER SPECIES MAZZAELLA SPLENDENS AND MAZZAELLA LINEARIS (RHODOPOHYTA)1

Phenotypic differences between the low wave exposure Mazzaella splendens (Setchell et Gardner) Hommersand and the high exposure Mazzaella linearis (Setchell et Gardner) Fredericq could be due to plasticity or genetic differentiation. Common gardens were used to assess their levels of plasticity after describing allometric relationships. As thalli lengthened, stipe development for M. splendens almost ceased even though blades continued to expand, but M. linearis formed a larger stipe before developing a blade that continued to stay narrow at longer thallus lengths. Common gardens demonstrated that M. splendens regrown in the site of M. linearis produced a wider blade than M. splendens regrown in its natural low energy site and that M. linearis regrown in low wave energy either could not form a wider blade or became narrower than thalli from its high energy site. Tetrasporophytes of M. splendens produced a longer and thicker stipe in the high energy site, but the larger M. linearis stipe was not realized because its wider blades made it vulnerable to hydrodynamic removal. Mazzaella splendens therefore had low survivorship in the high wave energy site, and survivors were not long enough to reproduce. Survivorship and reproduction of M. linearis was similar in both environments. Some of the M. splendens and M. linearis characters are plastic, but this plasticity was insufficient for convergence of phenotypes, and blade width plasticity was maladaptive at least for M. splendens. Developmental systems producing the stipe and blade phenotypes of each species have undergone genetic differentiation.     

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 genus Lessonia Bory (Phaeophyta,Laminariales) in Southern Chile and Argentina

Study of Lessonia in southern Chile and Argentina indicates that it is represented there by three species: L. nigrescens Bory, L. flavicans Bory and L. vadosa Searles sp. nov. Plants of these three species have been confused with each other in the literature, but appear clearly separable. Lessonia nigrescens is distinguished from the others by its massive, solid holdfast and numerous stipes. In Lessonia flavicans and L. vadosa a single stipe arises from a branched holdfast. These two species are distinguishable from each other in details of blade morphology, anatomy, and ecology. The taxonomic status of plants in South America outside of the study area remains unresolved. Populations in the Falkland Islands and in northern Chile in particular are in need of further study.     

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.     

TRANSLOCATION OF PHOTOSYNTHATE IN THE BROWN ALGA NEREOCYSTIS

Translocation of photosynthates was found to occur when blades of Nereocystis were illuminated in the presence of C¹⁴ bicarbonates. Rates of translocation averaging 37 cm/hr in the laboratory were observed. Samples from the epidermis, cortex, and medulla of the stipes of plants with blades exposed to tracer showed that the radioactivity in organic compounds was confined to medulla where sieve filaments occurred. Girdle preparations of blades, interrupting the mucilage ducts and leaving the blade medulla intact, allowed translocation to take place. These data indicate that conduction of photosynthate takes place in the medulla. Similarities between the anatomy of algal sieve filaments and angiosperm sieve tubes, coupled with the continuity of the sieve filaments from blade medulla to stipe medulla suggested indirectly that the sieve filaments were conducting elements. Further support of this hypothesis was provided from collections of radioactive exudate from cells in the medulla of the lower stipe that were continuous with the sieve filaments. Tracer applied to the blades was partially recovered as organic material in a clear fluid that collected inside the pneumatocyst. Continued accumulation of radioactivity in this fluid was dependent on living blades; fluid with low specific activity that did not increase during the experiment accumulated slowly if blades were killed with ethanol after an exposure to tracer. It is possible that the system that produced the stipe fluid was part of (or a side effect of) the system responsible for maintaining volume in the conducting system. It may also provide an alternate route (other than the sieve filaments) for delivery of photosynthates to the base of the plant. Carbon-14 applied to blade tips as bicarbonate was recovered in part as radioactive mannitol in the translocation stream.     

MORPHOLOGY AND MOLECULAR PHYLOGENY OF AUREOPHYCUS ALEUTICUS GEN. ET SP. NOV. (LAMINARIALES,PHAEOPHYCEAE) FROM THE ALEUTIAN ISLANDS1

A previously unknown species of kelp was collected on Kagamil Island, Aleutian Islands. The species can be easily distinguished from any known laminarialean alga: the erect sporophytic thallus is composed of a thin lanceolate blade attaining ～2 m in height and ～0.50 m in width, without midrib, and the edge of the blade at the transition zone is thickened to form a V‐shape; the stipe is solid and flattened, slightly translucent, attaining ～1 m in length; the holdfast is semidiscoidal and up to 0.15 m in diameter. Anatomically, the blade has the typical trumpet‐shaped hyphae characteristic of the Chordaceae and derived foliose laminarialean species (i.e., Alariaceae/Laminariaceae/Lessoniaceae). No hair pits or mucilaginous structures were observed on the blade or stipe. No fertile sporophytes were collected, but abundant juvenile sporophytes were observed in the field. In the molecular phylogenetic analyses using chloroplast rbcL gene, nuclear ITS1‐5.8S‐ITS2 rDNA, and mitochondria nad6 DNA sequences, the new species (Aureophycus aleuticus gen. et sp. nov.) showed a closer relationship with Alariaceae of conventional taxonomy, or the “Group 1” clade of Lane et al. (2006) including Alaria and related taxa than with other groups, although the species was not clearly included in the group. Aureophycus may be a key species in elucidating the evolution of the Alariaceae within the Laminariales. Because of the lack of information on reproductive organs and insufficient resolution of the molecular analyses, we refrain from assigning the new species to a family, but we place the new species in a new genus in the Laminariales.     

Field measurement of the dynamics of the bull kelp Durvillaeaantarctica (Chamisso) Heriot

Seaweed habitats and morphological development are strongly affected by wave forces. Novel measurements were made of the force dynamics of the large intertidal macroalga Durvillaeaantarctica under the influence of wave action. Synchronized video, a pressure sensor and a resistance wave gauge provided data describing the wave field. The response of seaweeds to waves was gauged using instrumentation mounted directly on the seaweed, including accelerometers and displacement and force transducers. These field measurements were used to estimate forces and bending moments acting at the holdfast, where failure is most likely to occur. For waves of the order of 0.5 m high, we measured maximum forces on the stipe of around 300 N and blade accelerations that exceeded 30 m s⁻². During large wave events, inferred bending moments at the base of the stipe reached average values of around 140 N m. There was a decoupling between the blade response and the force experienced at the stipe base. Furthermore, changes in water depth throughout the tidal cycle had a systematic effect on blade accelerations and moments at the holdfast.     

TRANSLOCATION OF 14C IN MACROCYSTIS INTEGRIFOLIA (PHAEOPHYCEAE)1,2

Translocation patterns in the giant kelp, Macrocystis integrifolia Bory, were investigated in situ using ¹⁴C tracer; sources and sinks were identified. Export was first detected after 4 h of labeling; experiments were routinely 24 h continuous ¹⁴C application. Mature blades exported ¹⁴C to young blades on the same frond and on younger fronds, as well as to sporophylls and frond initials at the bases of the fronds. Blades <0.3 m from the apex imported and did not export; this distance did not change seasonally. In spring export from blades 0.3–1.25 m from the apex was exclusively upwards; older blades also exported downwards. In fall downward export began 0.5 m from the apex, and blades >2 m from the apex exported exclusively downwards. Carbon imported by frond initials, young fronds, and sporophylls in fall may partly be stored for growth in early spring. No translocation was seen in very young plants until one blade (secondary frond initial) bad been freed from the apical blade; this blade exported to the apical blade for a time, but imported when it began to develop into a frond. The second and third formed blades on the primary fronds (sporophylls also exported when <0.3 m from the apex, and later stopped. Frond initials and sporophylls on later-formed fronds did not export at all. The translocation pattern in M. integrifolia differs from that previously reported in M. pyrifera in seasonal change and in distances from the apex at which the changes take place.     

Ecological effects of harvesting Lessonia (Laminariales,Phaeophyta) in central Chile

Lessonia nigrescens and L. trabeculata are economically important canopy-forming kelps in Chile. Experimental harvesting of stipes above the first dichotomy reduces stipe movement and inter-stipe friction, allowing the development of a heavy epiphytic load and increased grazing. Complete stipe removal leads to holdfast death as neither species is able to simultaneously regenerate all stipes. The invertebrate fauna inside the holdfast does not respond to upper canopy changes, but mortality does occur in partial or complete plant removals. Kelp removal also affects inter-plant distances, results in increased access of grazers to the outside and inside of kelp holdfasts, reduces recruitment of other algal species, and modifies the morphology of L. trabeculata such that the plants become more susceptible to removal by water movement.     

Growth and reproductive patterns of <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> sporophytes in a cultivation farm in Busan,Korea

Monthly growth and reproduction of Undaria pinnatifida sporophytes were examined over a period of 5 months in a cultivation farm in Korea. A total of 11 characters of Undaria were measured to determine a reliable morphological character representing its growth and reproduction. Plant weight of Undaria sporophytes increased steadily over the experimental period, but it increased in four different ways. Undaria pinnatifida increased body weight by growth in length and width (October–early December), and by growth in width with the thickening of blade and stipe when sporophytes began to be fertile (December–January). In the middle of January, growth in length and width had almost stopped with the maturation of Undaria sporophytes. Finally, the weight of Undaria increased again by growth in width at the end of February. Present results indicate that Undaria sporophytes increase body weight by growth in length and width at different times, and the relationship between reproduction and vegetative growth is exclusive. Plant weight was positively correlated and fitted well with stipe width and blade width. The blade of Undaria was very thin (ca. 254 μm) and breakable by wave action, but its stipe was strong and relatively thick (ca. 8.7 mm). Furthermore, the fertility of U. pinnatifida was fitted better with stipe width than blade width. Thus, we suggest that the stipe width is the most feasible character with which to estimate the growth and reproduction of U. pinnatifida sporophytes in the cultivation farm.     

LATITUDINAL TRENDS IN THE GROWTH AND REPRODUCTIVE SEASONALITY OF DELESSERIA SANGUINEA,MEMBRANOPTERA ALATA,AND PHYCODRYS RUBENS (RHODOPHYTA)1

The seasonality of Delesseria sanguinea, Membranoptera alata, and Phycodrys rubens (Rhodophyta) was studied at Helgoland (North Sea, Germany) and Roscoff (Brittany, France). Plants were collected at bimonthly intervals, and growth and reproduction were monitored. Growth of blades was observed mainly in spring, although small blades were found on plants of M. alata and P. rubens all year round. In summer, plants started to degenerate and in autumn they became fertile. The reproductive season of D. sanguinea lasted from October to February/April at both locations, whereas reproductive plants of M. alata and P. rubens were found until April at Helgoland and until June and August, respectively, in Roscoff_: Lower winter temperatures at Helgoland than at Rascoff may have caused these differences in the duration of the reproductive season. Using published data, the seasonal patterns at Helgoland and Roscoff were compared to those found at other locations (e.g. Barents Sea; Maine, USA; Isle of Man, UK) and local temperature/daylength conditions. Blade growth was synchronized across all populations and occurred in spring, when temperatures were usually still suboptimal for growth. Maximum reproduction was generally found in the colder half of the year but started earlier in autumn in the Barents Sea. Adaptive strategies in the seasonal control of growth and reproduction are discussed. Adequate timing of the history events (e.g. appearance of juveniles in spring) appears more important than maximal growth and reproduction of adults during the season with the most favorable temperatures.     

Microscopic observations on tissue calcium distribution in the brown alga,Undaria pinnatifida (Harvey) Suringar

The distribution of Ca²⁺ in thallus tissues of the brown algaUndaria pinnatifida was investigated by microspectro-photometric analysis after fixation with Carnoy's reagent and staining with an alizarine sulfonate dye. Ca²⁺ accumulation in the young thallus was much higher in mucilage cells of the basal than the apical region; there was least in the pith of the basal region. A roughly uniform pattern was found in the stipe from the central part of the young thallus, while a highly irregular pattern occurred in both stipe and blade from the corresponding area of the mature thallus; the Ca²⁺ content was much lower than in the cortex, mucilage cells and vacuoles of the mature stipe than in the corresponding area of the young stipe. The contents of all regions of the mature blade were higher than those in the young blade; they were particularly high in mucilage cells.     

Microbial populations associated with the surface of the brown algaAscophyllum nodosum

The microorganisms on the surface of the brown algaAscophyllum nodosum, collected from an intertidal area in Nahant, Massachusetts, were examined using scanning electron microscopy. Differences in the microbial populations on the holdfast, internodal regions of the stipe, and the apical tips were apparent. The populations ranged from a lawn of end-attached bacteria above the holdfast to microcolonies of yeast cells near the apical tips. The greatest diversity of microorganisms was noted in the internodal region representing the fourth year of growth where a dense lawn of end-attached bacteria was overlaid by filamentous bacteria, pennate diatoms, and filamentous blue-green algae. A simple procedure was developed to estimate the number of bacteria on the surface of the seaweed using the scanning electron microscope. The observed distribution of epiphytes may be explained in terms of the age of the algal surface, differences in light intensity, and the differential secretion of tannin by various parts ofAscophyllum.     

CYTOPLASMIC ORGANIZATION AND RHYTHMIC STREAMING IN GROWING BLADES OF CAULERPA PROLIFERA

Light- and electron-microscopic studies of the growing blades and their meristematic tips in Caulerpa prolifera have been correlated with time-lapse photographic studies. The growing blade may be divided into three zones based on the level of maturation. A “meristemplasm” is present at the tip of the growing blade and at sites of wounding. The cytoplasm of these growing regions has an abundant endoplasmic reticulum, Golgi bodies, amyloplasts, and nuclei, but lacks chloroplasts and a central vacuole. An intermediate zone lies between the white meristemplasm and the green, mature basal zone. The basal zone contains a parietal cytoplasm with organelles typical of the Chlorophyta and a dominant central vacuole. Two distinct systems of cytoplasmic streams occur in the basal zone and both contain packets of microtubules orientated parallel to the axis of the streams. As the blade matures and growth ceases, the dominant central vacuole forms up to the tip. In developing blades the tips become white in the absence of light as a result of basipetal movement of the chloroplasts. In plants kept on a 12-hr L: 12-hr D cycle, blade growth and chloroplast migration at the blade tip are rhythmic, and the peak occurs about 2-4 hr after initiation of the dark phase. Experiments are reported using continuous light or darkness after three 24-hr periods of 12-hr L: 12-hr D phasing.