A COMPARATIVE STUDY OF CELL DIVISION PATTERNS DURING GERMINATION OF SPORES OF ANEMIA,LYGODIUM AND MOHRIA (SCHIZAEACEAE)

Cell division patterns during germination of spores of Anemia (A. hirsuta, A. munchii, A. phyllitidis), Lygodium (L. circinatum, L. flexuosum, L. japonicum, L. salicifolium) and Mohria caffrorum have been examined by light microscopy of glycol methacrylate embedded materials. Spores of all species in a genus exhibited a constant pattern of division under different conditions of germination. In spores of species of Anemia, following an asymmetrical division, the proximal cell differentiated into the protonemal cell while the distal cell divided to produce the rhizoid. A similar division sequence was found in spores of Mohria caffrorum, but the fate of cells formed was reversed. In Lygodium spores, a proximal cell formed by an initial division of the spore cut off a protonemal cell, a rhizoid and a wedge-shaped cell by walls parallel to the polar axis. Our results contradict earlier observations on cell division sequence during germination of spores of these genera based on whole mount preparations.     

GROWTH REGULATION BY ETHYLENE IN FERN GAMETOPHYTES. V. ETHYLENE AND THE EARLY EVENTS OF SPORE GERMINATION

Early events during the germination of spores of the fern Onoclea sensibilis were studied to determine the time during germination when ethylene had its greatest inhibiting effect. Water imbibition by dry spores was rapid and did not appear to be inhibited by ethylene. During normal germination DNA synthesis occurred about four hours before the nucleus moved from a central position to the spore periphery. Following nuclear movement, mitosis and cell division occurred, partitioning the spore into a small rhizoid cell and a large protonemal cell. Cell division was complete approximately six hours after nuclear movement. Ethylene treatment of the spores blocked DNA synthesis, nuclear movement, and cell division. The earliest DNA replication in uninhibited spores was observed after 14 hours of germination, and the maximal rate of spore labeling with ³H-thymidine was between 16 and 20 hours. Spores were most sensitive to ethylene, however, during the stages of germination prior to DNA synthesis, and it was concluded that ethylene did not directly inhibit DNA replication but blocked germination at some earlier fundamental step. The effects of ethylene were reversible. since complete recovery from inhibition of germination was possible if ethylene was released and the spores were kept in light. Recovery was much slower in darkness. It was hypothesized that light acted photosynthetically to overcome the ethylene inhibition of germination. Consistent with this, it was shown that spores exhibit net photosynthesis after only two hours of germination.     

THE SPORE-GERMINATION PATTERN OF THELYPTEROID FERNS

Cell division patterns in germinating spores of several Thelypteris species were studied using light microscopy of sectioned material and scanning electron microscopy. All species exhibited the same basic germination pattern, characterized by an asymmetric cell division of the spore parallel to the equatorial plane to delimit a proximal rhizoid, followed by a perpendicular division of the basal cell to form the protonemal cell. While spore-germination patterns appear to be a potentially useful taxonomic character in some groups of ferns, the homogeneity in this character exhibited by the thelypteroid group impairs its usefulness in the taxonomy of Thelypteris.     

Analysis of the multi-stem clump structure ofLitsea japonica Juss. growing in a coastal dwarf forest

The multi-stem clump structure of a coastal dwarf forest dominated byLitsea japonica Juss. was investigated in order to clarify the sprouting characteristics and self-maintenance of clumps by stem alternation. The size and age distribution of multi-stem clumps were analyzed using cumulative relative frequency curves.L. japonica had a large number of stems and an even height distribution or young age-biased distribution of stems within a clump. These results indicated the sequential flushing of sprouts at high frequency. Height distribution within a clump ofL. japonica was relatively even compared to other species. This clump structure suggested the stable self-maintenance of individuals in all ranges of size and age without disturbances. It originated specific sprouting characteristics as a response to the severe stress of salty wind.Ardisia sieboldii Miq. had few stems within a clump. Although the stem height distribution of large individuals tended to be even, most clumps had a large size-biased distribution of stem height which indicated simultaneous sprouting. From this structure, sprouts of this species were thought to be of less significance in the stable self-maintenance of individuals thanL. Japonica.     

Spore Germination and Rhizoid Differentiation in Onoclea sensibilis: A Two-Dimensional Electrophoretic Analysis of the Extant Soluble Proteins

Following a geometrically asymmetrical cell division during germination of spores of the fern Onoclea sensibilis L., the small cell differentiates into a rhizoid and the large cell divides to form the protonema. Using silver-staining of two-dimensional gels, we have examined the soluble proteins of spores during germination and of separated rhizoid protoplasts and protonemal cells. Of over 500 polypeptides followed, nearly 25% increased or decreased in prominence during spore germination and the initial phases of rhizoid elongation. Soluble proteins from purified protoplasts of young rhizoids were quantitatively different from those of protonemal cells and germinated spores. Nine polypeptides which appeared after cell division were substantially more prominent in rhizoid protoplasts than in whole germinated spores and have been putatively designated rhizoid-specific polypeptides. The differences in the soluble protein composition of young rhizoids and protonemal cells probably reflect the differential organelle distribution between the two cells as well as differential net protein synthesis in the cytoplasms of the two cells.     

Growth strategy of the stoloniferous rattan Calamus javensis in Mt. Halimun, Java

Calamus javensis is a widespread understorey rattan in Southeast Asia and is one of the dominant rattan species in Mt. Halimun National Park, West Java. This species can establish clumps either sexually with seeds or clonality with stolons. To analyze the role of sexual reproduction and clonality for clump establishment in C. javensis, we studied the vegetative and genetic structures of the population. Totals of 2,777 ramets comprising 1,321 clumps were found in three 0.16 ha plots. The number of ramets per clump showed reverse J-shaped distribution with a maximum of 15. Clustering and stoloniferous ramets accounted for 62 and 20% of all ramets, respectively. This indicates that C. javensis is actively engaged in clustering clonal growth. The longest stem was 14 m although 75% of all stems were shorter than 1 m long. The stolon ranged from 6 to 538 cm in length, which decreased the local density of ramets. Using randomly amplified polymorphic DNA (RAPD) analysis, we obtained 56 polymorphic bands for 118 clumps sampled from one of the plots. All samples showed different RAPD banding patterns, except one pair at a distance of 225 cm from each other. The genetic similarity among samples closer than 10 m was significantly higher than the expected value from a purely random distribution. These results suggest that most separate clumps are established by recruitment from seeds, and most seeds dispersed near mother plants. Hence, in C. javensis, clonality is more a growth strategy to increase the size of the genet than a dispersion-propagation strategy to expand the habitat.     

Pattern and Performance of Understory Bamboos (Chusquea spp.) under Different Canopy Closures in Old-Growth Oak Forests in Costa Rica1

The effect of canopy closure on spatial and morphological patterns of three understory Chusquea species was studied in pristine upper montane oak forests. Plots with either Chusquea talamancenis Widmer & L.G. Clark or C. tomentosa Widmer & L.G. Clark were set out in gaps, under intermediate tree canopy and under closed canopy. The third species Chusquea foliosa L.G. Clark was only found in gap and intermediate canopy sites in the study area. Clumps of all bamboo species tend to be few and large under open conditions and numerous and small under more shady conditions. The larger size of clumps in gaps is reflected in the number of culms per clump, the diameter of the culms and, as a consequence, clump basal area, clump crown area, and biomass (index of plant volume) compared to intermediate and closed canopy. The smaller clump density in gaps compared to closed canopy implies that there is more intraspecific competition and density dependent mortality (self-thinning effect) when a bamboo species is dominant under favourable light conditions. Parameters of performance like culm length, number of nodes per culm, number and length of primary branches, and number of branch nodes seem not to be affected by the light regime, unlike total number of branches, total branch length, and the branching pattern. Species differ in their response to the light environment: C. tomentosa and C. foliosa have a higher degree of morphological plasticity than C. talamancensis, which in turn, appears to be more shade tolerant. No difference between species has been found regarding their contribution to vegetation, parameters reflecting abundance per unit area (site clump area, site crown area and site index of clump volume) were similar for all three species.     

Gametophyte-sporophyte coalescence in populations of the intertidal carrageenophyte <Emphasis Type="Italic">Mazzaella laminarioides</Emphasis> (Rhodophyta)

Mazzaella laminarioides has consistently been reported as a typical coalescent/clump species with a triphasic life history of the Polysiphonia-type in which the haploid gametophyte is the predominant phase with respect to the diploid sporophyte. Preliminary observations of intertidal populations revealed that, in some instances, cystocarpic and tetrasporic fronds emerged from the same clump (G-T clumps), implying a coalescent process of haploid and diploid thalli by fusion of their corresponding adjacent basal holdfasts. Population surveys at three sites in Coliumo Bay, central Chile, were carried out to characterize frond demography as well as to asses the frequency of gametophyte-tetrasporophyte (G-T) coalescence. Visual and resorcinol methods were employed to determine the phases of the fronds collected over central transects of 15 randomly sampled clumps. Coalescence of G-T clumps was infrequent, with gametophytes dominating over tetrasporophyte thalli.     

Establishment of the Woody Grass Arundinaria gigantea for Riparian Restoration

Canebrakes are dense stands of Arundinaria gigantea (Walt.) Muhl. that covered large areas of the southeastern North America. With agricultural development, canebrakes were quickly converted to crop and pastureland and now occur only in small, isolated patches. There is growing interest in the use of A. gigantea and other temperate bamboo species in riparian and floodplain revegetation in North America, but lack of detailed information on propagation and management of woody perennial grasses hinders reestablishment activities. Our study assesses the influence of nutrient and woodchip mulch amendments on survival and growth of A. gigantea transplanted as part of a riparian restoration project in central Kentucky. After two growing seasons, culm number (aboveground stems) increased 4‐fold and extent of transplanted clumps expanded 26‐fold. The survival rate of transplanted cane clumps was 98%. Hardwood chip mulch significantly increased the emergence of new culms, culm height growth, and clump area. Composted manure, applied at a rate that contributed a similar mass of organic matter as the hardwood mulch, also significantly increased new culm number and clump area. Our findings demonstrate that addition of manure or hardwood mulch can significantly enhance aboveground production of A. gigantea transplants. However, survival and initial growth of untreated clumps were also adequate in this study. It appears that careful site selection, transplantation, and site maintenance may be sufficient to ensure A. gigantea establishment on many sites. Practitioners should assess soil drainage, water stress, and fertility along with herbaceous competition and incidence of overbank flooding before determining the necessity of organic amendments to supplement establishment of A. gigantea or other woody grasses for riparian restoration.     

Spore wall morphology and dehiscence pattern in the liverwort Fossombronia caespitiformis De Not.

Abstract

The spore wall morphology and first germination stages of Fossombronia caespitiformis are described. The spores are markedly heteropolar and catalept. The boundary line, which separates the proximal and distal faces, is shifted toward the proximal pole. The spore coat breaks along this boundary line allowing the protonemal tube to protrude. The dehiscence pattern can be considered tangential.     

Bamboo,fire and flood: consequences of disturbance for the vegetative growth of a clumping,clonal plant

The clumping growth form of tropical bamboos suggests a consolidator strategy in the face of intense competition rather than an ability to exploit disturbance. We investigated the annual growth and culm demographic responses to disturbance by fire and flood of vegetatively mature clumps of a riparian stand of Bambusa arnhemica. Linear-mixed models were employed to control for the non-independence of culm fates within clumps and clump growth among years. As the stand was monodominant and the species is gregariously monocarpic, responses can be interpreted independently of interspecific competition and trade-offs with sexual reproduction. Disturbance depressed clump growth but the culm demographic responses to fire and flood were quite different. Few culms were destroyed during the fire but damage depressed their subsequent survival, leading to declines in clump basal area. Fire also triggered the release of ramet buds, but the additional recruits were small and short-lived. Prolonged and early flooding in one of the study years depressed culm recruitment in clumps low on the lagoon bank but there was possible compensatory recruitment in the following year. There was no convincing evidence that sparse clumps recruited culms better than dense clumps, though culm recruitment was negatively correlated with culm survival. Fire and prolonged flooding are inevitable elements of the environment occupied by B. arnhemica, but we interpret the species’ response as persisting in the face of disturbance rather than exploiting the opportunities created by it.     

On the benefits of living in clumps: a case study on Polytrichastrum formosum

• The study concerns the mechanics and water relationships of clumps of a species of endohydric moss, Polytrichastrum formosum.
• Anatomical and morphological studies were done using optical and scanning electron microscopy. Experiments on waterdrop capture and their distribution to adjacent shoots within a moss clump were performed with the experimental set‐up for the droplet collision phenomena and ultra‐high speed camera. The mechanical strength of the moss clump was tested on an electromechanical testing machine.
• During the process of moss clump wetting, the falling water drops were captured by the apical stem part or leaves, then flowed down while adhering to the gametophore and never lost their surface continuity. In places of contact with another leaf, the water drop stops there and joins the leaves, enabling their hydration. Mathematical analysis of anatomical images showed that moss stems have different zones with varying cell lumen and cell wall/cell radius ratios, suggesting the occurrence of a periodic component structure. Our study provides evidence that the reaction of mosses to mechanical forces depends on the size of the clump, and that small groups are clearly stronger than larger groups.
• The clump structure of mosses acts as a net for falling rain droplets. Clumps of Polytrichastrum having overlapping leaves, at the time of loading formed a structure similar to a lattice. The observed reaction of mosses to mechanical forces indicates that this phenomenon appears to be analogous to the ‘size effect on structural strength’ that is of great importance for various fields of engineering.

     

CONVERGENT BIOLOGICAL PROCESSES IN COALESCING RHODOPHYTA

Sporeling coalescence in Gracilaria chilensis Bird, McLachlan et Oliveira produces genetically polymorphic, chimeric individuals. If this is common in red algae, it may have significant biological consequences. In this study, we evaluate the hypotheses that coalescence is widespread among the Rhodophyta and that specific and convergent morphological and ecological responses characterize this as a unique growth style among marine algae. A literature survey on coalescence was undertaken to assess the distribution of this condition in the Florideophycidae. Sixty-two (54.9%) of 113 species considered germinated to form a disk. Subsequent development in 37 of these species showed crust formation and coalescence during development with other crusts in 31 species (84%). Coalescing red algae were members of the orders Ahnfeltiales, Corallinales, Gigartinales, Gracilariales, Halymeniales, Palmariales, and Rhodymeniales. Ultrastructural studies in species of Ahnfeltiopsis, Chondrus, Gracilaria, Mazzaella, and Sarcothalia suggested a common pattern of early development. Newly released, naked spores may fuse into a single cell, as they do in Chondrus canaliculatus, or they may develop individual cell walls that later are surrounded by a thickened common wall. Ultrastructural studies demonstrated two kinds of immediate development after the first mitotic division: direct development by symmetric divisions resulting in discoid sporelings or an indirect asymmetric arrangement of divisions before a diskoid sporeling was formed. Germination in coalescing species is a linear function of the initial spore density, whereas in noncoalescing species maximum germination occurs at intermediate densities. In the field, coalescing species may recruit either from solitary or aggregated spores. However, survival is significantly higher for plantlets grown from a larger number of coalescing spores. Total number of erect axes formed by the coalesced mass is a logarithmic function of the initial number of spores. Thus, germlings grown from a larger number of coalescing spores exhibited a larger photosynthetic canopy than do plantlets grown from a few spores. Juveniles and mature clumps grown from a coalescing mass may exhibit size inequalities among erect axes, with the larger axes located toward the center of the clump. These larger axes mature first or, in some cases, are the only to produce spores. The widespread occurrence of coalescence inroughly half the number of orders of the Florideophycidae, the similarity of the coalescence process, and the finding of various adaptive traits associated with coalescence characterizes this as a unique growth style, splitting the diversity of species now included in the Florideophycidae into two major groups: coalescing and noncoalescing Rhodophyta.     

Funaria Rhizoids infected with Pleotrachelus wildermani

Abstract

The recolonisation of bryophytes on soil during 3½ years following fire in a mixed forest in south-west Tasmania is characterised by the presence of Marchantia berteroana, Funaria hygrometrica, Ceratodon purpureus and Polytrichum juniperinum, while limited cover is provided by Campy lop us introjlexus, Tortella calycina and Bryum sauteri. It has been verified that the protonemal growth of the early colonising mosses is more rapid than non-colonising specles. The preference for spores of Funaria hygrometrica, Ceratodon purpureus and Marchantia berteroana to germinate on burnt soil is confirmed. However, it has been shown that burnt substrates do not inhibit the germination of non-colonising species. Therefore, the rapid development of protonemata and gametophytes in the colonising species may be a critical factor for their success.     

Sex ratio from germination through maturity and its reproductive consequences in the liverwort Sphaerocarpos texanus

Summary The dioecious winter ephemeral liverwort, Sphaerocarpos texanus disperses spore tetrads consisting of two males and two females. I examined the subsequent sex ratio of S. texanus at different stages in its life cycle to detect possible mechanisms affecting deviations from a 1:1 sex ratio and the effect of sex ratio on reproductive success. As S. texanus occurs in pure male, pure female and mixed sex clumps, I examined the proportions and sizes of these, the reproductive success of pure female and mixed sex clumps in the field and the sex ratio of germinating plants in a growth chamber. In both the field and the growth chamber the most abundant clump type was pure female followed by mixed sex and pure male clumps. These abundance patterns suggest that males have a lower survival rate than females before emergence and this lower survival rate continues through the gametophytic stage. This disadvantage may be due to the higher susceptibility of males to environmental conditions, to their competitive inferiority to females, and/or to differential resource allocation to the sexes within the spore tetrad. The female biased sex ratio at germination is consistent with predictions from sex ratio theory. Further my field data indicate that males may gain a survival benefit from growing in a mixed sex clump and both males and females benefit reproductively when they occur in mixed sex clumps.     

A stable inhibitor of spore germination produced by fungi

Summary The level of inhibition of germination of spores added to cell-free culture filtrates of Fusarium oxysporum Schlecht. ex Fr. declines with time. This decline can be accounted for by the decreasing levels of volatile sporostatic factors and a stable vacuolation factor produced by F. oxysporum. After the disappearance of the volatile sporostatic factors, a stable principle is recognised in the culture filtrate by a persistent level of inhibition of spore germination which is not completely alleviated by the addition of nutrients. A stable inhibitor of spore germination has been isolated and characterised as nonanoic acid, and evidence is presented that this compound is widely produced among fungi and can inhibit the germination of spores of many fungal species. The contribution of nonanoic acid to sporostasis in cultures of F. oxysporum is assessed.     

DNA Synthesis and Cell Division During Spore Germination in Lygodium japonicum

This paper describes the ontogenetic sequence of cell divisionsand associated DNA synthetic patterns observed in sectionedspores of Lygodium japonicum (Thunb.) Sw., collected at differentstages of germination. Following exposure to a saturating doseof red light, the spore undergoes an asymmetric division toform a basal cell, which retains nearly all of the storage inclusions,and an apical cell which expands and protrudes from the rupturedsporoderm. Division of the apical cell results in formationof a protonemal cell and an intermediate cell. Subsequently,the latter cell divides to form the primary rhizoid and a wedgecell adjacent to the protonemal cell. Secondary rhizoids mayarise from later divisions of either the basal cell or the wedgecell. In addition, the wedge cell appears to have the capacityto form a secondary prothal-lial filament. Histochemical localizationof cell constituents indicates an increasing concentration ofcytoplasmic RNA and protein in the presumptive protonemal regionof the spore cell prior to division. Autoradiography of ³H–thymidineincorporation has shown that synthesis of nuclear DNA precedeseach cell division. Although strictly nuclear DNA synthesisoccurs during early stages of germination, extra-nuclear DNAsynthesis increases greatly following division of the sporecell. The results are discussed in relation to earlier studieson cell division patterns seen in whole mount preparations ofgerminating spores of different species of Lygodium. Lygodium japonicum, spore germination, cell division, DNA synthesis     

Colonization of floating seaweed by pelagic and subtidal benthic animals in southwestern Iceland

Coastal floating seaweed, originating from the intertidal, is colonized by various pelagic and subtidal benthic animals. At southwestern Iceland, common macrofaunal colonizers include the fish Cyclopterus lumpus L., the harpacticoid Parathalestris croni (Krøyer), the amphipods Calliopius laeviusculus (Krøyer), Gammarellus angulosus(Rathke), Dexamine thea Boeck and Ischyrocerus anguipes (Krøyer), and the isopod Idotea baltica Pallas. The colonization patterns of these species were studied by collecting samples from experimental and natural clumps of floating seaweed at approximately monthly intervals for ca. 18 months. The occurrence of colonizers was highly seasonal with maximum numbers between April and September. For most species, numbers were correlated with weight of clumps. The relationship of density (numbers/100 g algae) to weight of clump and distance from shore varied between species. The algal species composition of clumps appeared to influence some colonizers. Only juvenile C. lumpus were found, while all P. croni were sexually mature; juveniles predominated in other species. The different colonization patterns of the species can be explained partly by their biological traits. The clumps are vital for the breeding of P. croni and are used for feeding by C. lumpus fry. Other species possibly colonize floating algae accidentally, being programmed to seek attached vegetation. Floating clumps may, nevertheless, serve to disperse these species.     

Effects of Multispecies Clumping on Survival of Three Succulent Plant Species Translocated onto Mine Spoil in the Succulent Karoo Desert, South Africa

Salvaging from premining areas and translocation of succulent plants has been investigated for the revegetation of gypsiferous mine spoil in an arid region of South Africa. Given that facilitation effects are thought to outweigh competition effects in harsh environments, we hypothesized that the survival of translocated succulents would be higher when planted in multispecies clumps than alone and that the growth rate (measured as stem extension) and fruit‐set would be greater for plants in clumps than for those planted alone. Two leaf‐succulent species (Aridaria noctiflora ssp. noctiflora and Drosanthemum deciduum) and one stem‐succulent species (Psilocaulon dinteri) were salvaged from the area destined for mining and translocated onto the mine spoil. These plants were planted either in a multispecies clump of the three species together or alone. One year after translocation, 67% of the plants survived. It was also found that the succulents used in these experiments survived in higher numbers when planted alone. Due to the similar root morphology of D. deciduum and P. dinteri, they competed for resources instead of facilitating each other's establishment. The results were variable for each of the species used, and neither growth nor seed‐set was improved by clumping. These findings would also possibly vary from year to year with different abiotic conditions.     

Multiple allelopathic activity of the crustose coralline algaLithophyllum yessoenseagainst settlement and germination of seaweed spores

A study was made to investigate possible formation by the crustose coralline algaLithophyllum yessoenseof multiple allelopathic-related substances against the settlement and germination of spores of various seaweeds. Seven different solvents (n-hexane, diethyl ether, acetone, ethyl acetate, acetonitrile, methanol, distilled water) and seawater were used to obtain crude extracts and secretory exudates from the coralline alga. The extracts and the algal conditioned seawater were tested for inhibitory activity against the settlement and germination of spores from 17 species representing 15 genera. Spore settlement of 14 species was inhibited over 90% by one or more extracts of the six organic solvents and conditioned seawater. The germination of spores from 13 species was inhibited by one or more extracts of all seven solvents and conditioned seawater. The species where spore settlement was not significantly affected showed strong inhibition of germination, andvice versa.