Architectural models for apical patterns in Gelidium (Gelidiales, Rhodophyta): hypothesis of growth

The great morphological variation in the apical region of main axes and branches in species of Gelidium and Pterocladia suggests different patterns of growth. Those patterns have been reconstructed from the ontogenetic sequence of morphological stages and regeneration of the apex. Using the architectural approach, models for four species of Gelidium from the Pacific coast of central Chile and the Pacific coast of Mexico's tropical region, were constructed based on the following criteria: growth pattern, specialization of laterals, symmetry, apical dominance and rhythm of apical and lateral growth. Those models allow for a formal evaluation of apical differences between species.     

Intrageneric differences in cystocarp structure in Gelidium and Pterocladia

Unilocular cystocarps, with ostioles opening to one frond surface, have traditionally distinguished Pterocladia from Gelidium, described as having bilocular cystocarps, with ostioles opening to both surfaces; however, unequally developed locules have been described in Pterocladia and differences in cystocarpic architecture between Pterocladia capillacea and the type species of the genus, P. lucida, have been recently found. As heterogeneity in cystocarp architecture raises questions of basic intergeneric distinction, a survey of reproductive morphology of species in both genera is presented in this study. Six morphologically-different types of cystocarps are distinguished among the five species of Pterocladia and the seven species of Gelidium examined.     

The Structure and Development of the Reproductive Organs and Carposporophyte in two British Species of Gelidium

The structure and development of the reproductive organs andcarposporophyte are described for two British species of Gelidium,G. latifolium (Grev.) Born. & Thur. and G. pulchellum (Turn.)Kütz. The gonimoblast does not develop directly from theunchanged carpogonium, as was thought by Kylin, but a swollenmultinucleate cell of irregular outline is formed first, eitherfrom the carpogonium alone, or by the fusion of the carpogoniumand certain neighbouring cells; the gonimoblast develops fromlong, non-septate processes of this multi-nucleate cell. Thesignificance of the fusion of the carpogonium and certain vegetativecells is discussed in relation to the various definitions ofthe auxiliary cell which have been proposed. It is concludedthat an auxiliary cell does not occur in Gelidium.     

Medullary structure differences in Gelidium and Pterocladia (Gelidiales, Rhodophyceae): Taxonomic meaning

Study of the apical portions of vegetative thalli of 21 species of the genera Gelidium and Pterocladia (Gelidiales-Rhodophyta) revealed the existence of six patterns of medullary structure. Each pattern is characterized by the organization of the medullary tissue and the changes in cellular dimensions from apex to subapex. These characters can be used to separate groups of species within each genus.     

Ricerche Sulla Rigenerazione in Coltura Di Pterocladia Capillacea (Gmel.) Born. Et Thur

Abstract

Research on the regeneration in «Pterocladia capillacea» (Gmel.) Born, et Thur. cultured «in vitro». – It has been show that small segments (even 0,3 cm. long) of the thallus of «Pterocladia capillacea» undergo the regeneration «in vitro» of adventitious buds. Sea water enriched according to MIQUEL'S formula (1890) was used for the culture. Adventitious buds were present on both the cut-surfaces of the intercalary segments. Both surfaces regenerated, suggesting the absence of clear polarity for regeneration; howewer, usually one cut-surface showed more vigorous adventitious branches than the other one. The possibility that the predominance of one surface could be related to a polarization of the flow of nutrients from the parent segment has been discussed. The thallus originated «in vitro» is very similar to the creeping axes and the formation «in vitro» of thallus similar to that of the erect frond was never observed. 0,1γ/ml. IAA promoted regeneration and stimulated the growth of adventitious buds. IAA stimulated also the growth of the apical portion of the erect fronds. 10^?5M 2-chloro-ethyl trimethyl-ammonium chloride (CCC) stimulated regeneration and the growth of the tips of the main axis and lateral branches of apical segments. Higher CCC concentrations (10^?3M) inhibit regeneration and at the same time all the tips were necrotized; consequently some outgrowths appeared along the original axes. Such phenomenon was interpreted as a new type of regeneration obtained without any injury. Data reported in this preliminary communication represent a good approach for new study on the biochemical aspects of regeneration.     

THALLUS ORGANIZATION AND DEVELOPMENT IN THE FRUTICOSE LICHENASPICILIA CALIFORNICA,WITH COMPARISONS TO OTHER TAXA

Thallus organization is examined inAspicilia californicaRosentreter, a fruticose lichen known from several localities in central and southern California. The sprawling, terete thallus branches possess a dense central medulla of thick-walled, longitudinally oriented fungal cells. This central tissue emerges at branch apices to form a darkly pigmented fungal tip. Thallus development involves the apical extension of the tip to produce a fungal tissue over which a cylindrical algal layer and cortex will eventually be formed. Apical branches are initiated by furcation entirely within the fungal tip. Lateral branches, emerging from the lichenized thallus, arise as a divergent bundle of elongate fungal cells originating in the medulla. The photobiont appears to play no direct role in initiation of apical or lateral branches. It is concluded that thallus development inA. californicaoccurs with a relatively low degree of synchrony between mycobiont and photobiont growth, similar to the pattern observed in crustose lichens with prothallic growth. A rather similar type of thallus organization is observed inA. hispida, although in that species mycobiont growth and branch initiation appear to be somewhat more closely associated with algal cell proliferation. A squamuloseAspiciliafrom central Spain produces rhizomorphs that may sometimes become invested with an algal layer and cortex, resembling the thallus axes ofA. californica.     

Sur le développement des spores et la formation du thalle rampant de Kallymenia microphylla J. Ag. (Rhodophyceae,Cryptonemiales)

Abstract

On the spore development and the growth of Kallymenia microphylla J. Ag. (Rhodophyceae, Cryptonemiales) creeping thallus. — The filamentous creeping axis produced by spore grows into a terminal cell and then branches out into a monostratified creeping thallus looking like a disc. This creeping thallus is looking exactly like that of the Rhodocorton haucki (Schiffner) Hamel. Compared to the other species of Kallymenia, we find out that this creeping thallus has cellular fusions as well; they are more frequent and appear quite early here.     

Pattern of Shoot Growth in Zizyphus mauritiana and Z. oenoplia

Shoot polymorphism and patterns of sylleptic branching and shoottip abscission in two Zizyphus species are reported. Z. mauritiana(ber) produced three types of proleptic shoots: vigorous, normaland spur-type, by reiteration or following pruning;Z. oenopliahad only one type of shoot. Both species revealed up to thirdorder sylleptic branching on the vigorous proleptic shoots witha characteristic pattern and rhythm. Bud dormancy breaking chemicalsgiven as pre-pruning foliar sprays in ber, did not alter thepattern of sylleptic branching. Zizyphus oenoplia produced morefirst and second order sylleptic branches compared to Z. mauritiana.All main axes and 97% of first order sylleptic branches ofZ.mauritiana abscised their apical buds during summer, but inZ. oenoplia all main axes and 54% of first order sylleptic branchesremained active. Shoot tip abscission was almost complete onthe higher order sylleptic branches of both species. Activefirst order branches in Z. oenoplia were confined to the tophalf of the shoot. Both species had a few dormant apical budsduring the summer on their sylleptic branches, emerging mostlyfrom the middle portion of the shoot. Some of the most vigorousfirst order branches of Z. oenoplia, which had dormant apicalbuds during summer, showed a change in the frequency of syllepticbranching when they resumed growth following monsoon showers.These characteristic growth and branching patterns may haveadaptive value for canopy development under arid and semi-aridconditions. Copyright 1999 Annals of Botany Company Ber, branching pattern, shoot polymorphism, shoot tip abscission, sylleptic branching, Zizyphus mauritiana, Zizyphus oenoplia.     

Production of plantlets of the red alga of Gelidium genus (Rhodophyta) from thallus fragments

Experimental research on production of plantlents from fragments of thallus of the alga Gelidium were carried out. The best results on development of plantlets without rhizoids were achieved with cultivation of fragments of subapical parts of the plant. Thus, the average growth rate of plantlets in the last for 4 weeks of rearing was 6.7 ± 1.2% a day. Gelidium plantlets with rhizoids were grown from the top branches of plants in an intensively stirred culture with shading of the lower parts of the branches. It is suggested to use the same method for obtaining planting material during bottom rearing of Gelidium in the sea as that for intensive culture in tanks.     

New and old problems in the taxonomy of the Gelidiales (Rhodophyta)

The order Gelidiales includes over 140 agar-producing taxa. Many species are taxonomically confused; the boundaries of one family and all but four genera recently have been contested, and the controversy over ordinal status has lasted for over 25 years. This study reviews recent developments in the taxonomy of the group and suggests future areas for studies. The order cannot be defined by a few exclusive characters, as intended in the past, but it shows a unique combination of characters. Additional studies on Gelidiella and Acanthopeltis seem advisable to clarify family limits. The segregation of Onikusa and Pterocladiastrum is doubtful. Only three of six characters discriminate Gelidium from Pterocladia. None allows complete generic segregation and all need variability studies. Analysis of nomenclatural types in the light of morphological variation would permit an understanding of species limits in Gelidium and Pterocladia. Similar studies are needed in Gelidiella and Ptilophora.     

Sind Farne Kormophyten? Eine Alternative zur Telomtheorie

Modern ideas concerning cormophyte phylogeny are strongly influenced by the telome theory. Erect growth, radial symmetry, dichotomy, protostely and eusporangial structure are considered to be primitive features. Yet, the cormus needs redefinition. TheHypolepidaceae (s. str.) are shown to have rather a two-dimensional thallus than a three-dimensional cormus. Their so-called rhizome develops a continuous areophore margin which is connected with the margin of the fronds. The longitudinally inserted fronds and branches are produced by the terminal rhizome meristem which itself grows in many respects like the apical frond meristem. The rhizomes ofDennstaedtiaceae and theDavallia-type of rhizomes are intermediate between thisHypolepis-type and the true cormus-type. TheHypolepis-type is compared with fern prothalli. Under perhumid tropical conditions higher land plants may have evolved from small, creeping, two-dimensional fern prothallium-like progenitors with isomorphous gameto- and sporophytes.

Vorgetragen auf der Morphologentagung in Zürich 1975.     

Advances in cultivation of Gelidiales

Currently, Gelidium and Pterocladia (Gelidiales) are collected or harvested only from the sea. Despite several attempts to develop a cultivation technology for Gelidium, no successful methodology has yet been developed. Initial steps towards developmental efforts in Portugal, Spain, South Africa and Israel have been published. More developments have probably been performed but have not been published. Two different technological concepts have been tested for Gelidium cultivation: (1) the attachment of Gelidium fragments to concrete cylinders floating in the sea, and (2) free-floating pond cultivation technology. These vegetative cultivation technologies might be partially optimized by controlling physical, chemical and biological growth factors. The pond cultivation technology is the much more controllable option. The effects of all factors are discussed in detail in this review. It seems that the main difficulty with cultivation of Gelidium is its low growth rate. The claimed yields of the two technologies are far from being economically attractive at this stage of their development. It seems that in order to introduce Gelidium into commercial cultivation, major efforts in genetic improvement through selection or genetic engineering will be required. Only high yield strains will have the potential to compete economically with the present harvesting tradition. However, accumulated experience with genetic improvement of other useful seaweed species suggests that this is possible.     

Studies on the Development of the Air Pores and Air Chambers of Marchantia paleacea: 1. Light Microscopy

The ontogeny of the air pores and air chambers of Marchantiapaleacea begins with the schizogenous development of protodermalintercellular spaces of the initial apertures, and is completedwith the formation of the air pores and giant sub-epidermalair chambers bearing numerous photosynthetic filaments. Intercellularspace formation commences from the thallus surface and proceedsinwards to the first internal layer of cells. The cells amongwhich spaces develop do not originate from one mother cell.Spaces are formed only in the regions of the intersection ofthe anticlinal walls of three, four, or sometimes more successivederivatives of S₁, S₃ and S₄ segments of the apical cell, oneor two of which have been divided periclinally and the restanticlinally. Protodermal intercellular spaces appear in mostor all the corners of these cells, the anticlinal walls of whichexhibit an opposite disposition. The S₁, S₂, S₃ and S₄ segmentsare produced by definite divisions of a five-sided apical celland by a series of divisions give rise to initial cells of theinternal layers of the thallus and initial cells of the protodermaland sub-protodermal layers. The concept of a quiescent apicalcell cannot be accepted, since dividing apical cells have beenobserved, and the pattern of wall disposition of the thallusapex cannot be explained without the active participation ofthe apical cell. The air chambers are apparently of exogenous origin. They resultfrom the broadening of the bottom of the initial apertures bythe coordination of the rate of anticlinal divisions and growthof the protodermal and sub-protodermal cells surrounding theintercellular spaces of the initial apertures. The ontogenyof the pore rings starts at an advanced stage of air chamberformation not from a mother cell but from the cells which surroundthe closed entrance of the air chamber, by a shift of the planeof division from anticlinal to periclinal. Before the periclinaldivisions a new axis of growth perpendicular to the thallussurface is established in the mother cells of the pore. By a polarized growth into the air chamber followed by periclinaldivisions, the cells of the floor form initial cells of thephotosynthetic filaments. The latter divide again to form singleor branched photosynthetic filaments. Marchantia paleacea, air pore, air chamber     

Production ecology of Gelidium

Available data on determinants of production in species of Gelidium suggest several general patterns. Species diversity is higher in tropical latitudes, whereas in temperate latitudes the size of the fronds is larger, the species are ecologically dominant and commercially viable. Typically, the species occur on rocky substrate, often on coralline crusts, associated with rapid water movement and arranged in successive belts that can extend down to 25 m depth. Yields vary among species, to a maximum of 2.0 kg m^–2 y^–1. Growth and production in many species can best be explained by complex interactions between irradiance and nutrients. Temperature can interact synergistically with irradiance, while water movement compensates for nutrient limitations. Increased water movement or the addition of nutrients can prevent, to an extent, bleaching by high light and high temperature. Available data suggest the existence of at least eight biological factors affecting predicted productivity of Gelidium crops: morphology, age of the frond, thallus part, reproductive state, seasonality, crop density, life history phase and geographic and ecological origin of the species. At least four events can remove or destroy Gelidium crops: extreme low tides, storms, grazing and careless harvesting. Only the last-named factor has been analyzed over more extensive experimental periods.     

The Taxonomy of Sphacelaria cirrosa (Roth.) Ag., Sphacelaria fusca (Huds.) Ag., and Sphacelaria furcigera (Kutz.) Sauv: A Simple Statistical Approach

An attempt has been made to apply some simple statistical techniquesto the taxonomy of a problematical group of species within thegenus Sphacelaria Lyngb. Measurements were carried out on materialcollected from the field, material made available from Herbariumcollections and on material grown under experimental conditionsThe main characteristics considered are cell and filament dimensions,although certain reproductive structures have also been measured. It has been found that the number of longitudinal divisionsin each segment of the main axes and the primary branches hasa profound effect on the over-all appearance of the fronds ofthe three species investigated. Segment length breadth ratioand frond stiffness have been shown to be dependent on the numberof longitudinal walls. This characteristic has enabled one ofthe species, S. cirrosa (Roth) Ag., to be readily distinguishedfrom the other two. Another factor shown to be important is the determinate patternof growth displayed by S. cirrosa as opposed to the indeterminatepattern displayed by S. fusca (Huds ) Ag. and S. furcigera (Kutz.)Sauv. The pattern of growth affects the number of longitudinalwalls, filament width, and the relative lengths of the primarybranches. The angle at which branches emanate from the main axes has beenshown to be a useful characteristic in this group of species. The variability in the number of arms produced by each propagulepresents an interesting problem which has only been described,experimental work suggesting no explanation for this phenomenon.The degree of variability is clearly different in each speciesand is useful taxonomically if a large enough sample is availablefor examination.     

Artificial sporeling and field cultivation of Gelidium in China

The supply of Gelidium resources is dependent on collection from natural habitats. As these resources are very limited, one possible means of augmenting production is by implementation of artificial cultivation as has been done with Laminaria, Porphyra and Eucheuma. Chinese phycologists have conducted research on sporeling and field cultivation of Gelidium for many years. Trials have been made using: (1) raft cultivation based on vegetative propogation of thallus fragments, (2) cultivation based on spore-collecting and land-based tank culture to provide seedstock, (3) cultivation based on regeneration from small thallus fragments to provide seedstock. Some of these results have been promising, but development remains at the experimental stage and methods of cultivation need to be improved. This paper is an updated review of the research and development on artificial sporeling production and field cultivation of Gelidium in China.     

DICROGLOSSUM CRISPATULUM GEN. ET COMB. NOV. FROM WESTERN AUSTRALIA,REPRESENTING A NEW TRIBE WITHIN THE DELESSERIACEAE (RHODOPHYTA)1

Dicroglossum gen. nov. (Delesseriaceae, Ceramiales) is a monotypic genus based on Delesseria crispatula, a species originally described by Harvey for plants collected from southwestern Western Australia. Distinctive features of the new genus include exogenous indeterminate branches; growth by means of a single transversely dividing, apical cell; absence of intercalary divisions in the primary, secondary, and tertiary cell rows; lateral pericentral cells not transversely divided; not all cells of the secondary cell rows producing tertiary cells rows; all tertiary initials reaching the thallus margin; midrib present but lateral nerves absent; determinate lateral bladelets arising endogenously; blades monostromatic, except, at the midrib; carpogonial branches restricted to primary cell rows, on both surfaces of unmodified blades; procarps produced on both blade surfaces, each procarp consisting of a supporting cell that bears two four-celled carpogonial branches and one sterile-cell group of three to four cells; and tetrasporangia borne in two layers, separated by a central row of sterile cells. The combination of exogenous indeterminate branching and bicarpogonial procarps is considered to warrant the recognition of a new tribe, the Dicroglosseae, within the subfamily Delesserioideae.     

MORPHOLOGY OF NECTRIA HAEMATOCOCCA

Ascocarp development in Nectria haematoccocca begins with the formation of deeply staining coils as lateral branches of the vegetative hyphae. As these coils develop into multicellular, multi-nucleate ascogonia, they are surrounded by a pseudoparenchymatous envelope. During ascocarp development an apical meristem produces cells that elongate downward into the centrum, forming long, filamentous, apical paraphyses. When fully developed the cells of the apical paraphyses swell, producing a tissue that is pseudoparenchymatous in appearance. The ascogonium proliferates to form a layer of multinucleate ascogenous cells across the base of the ascocarp. Asci form from the ascogenous cells by means of croziers. The asci grow up among the apical paraphyses, which disintegrate as the ascocarp matures. This pattern is typical of the Nectria-type of development, indicating that this species belongs in the Hypocreales.     

Morphogenetic effects of daylength in Schottera nicaeensis

The S. nicaeensis thallus exhibits a typical secondary heterotrichy: terete perennant creeping axes and flattened annual erect fronds. The prostrate system produces new uprights in autumn, while in late spring the old blades produce marginal terete proliferations, which usually attach themselves to the substratum becoming new creeping axes. The present study demonstrates that the changes in morphogenetic trends are controlled by daylength, regardless of temperature. Both the upright formation and the flattened growth are short day responses, whereas both the terete proliferation outgrowth and the rhizoid production are long day responses. This seems to be the first report on two vegetative photoperiodic responses to different daylengths within a single algal species.     

Daylength and development in four species of Ceramiaceae (Rhodophyta)

Developmental patterns in four species of Ceramiaceae were determined using excised thallus apices grown under a range of light periods. Models of thallus development and organization based on these patterns are presented. Increased rates of apical cell division, greater growth of apical fragments and increased average cell size were found with increasing number of hours light per day between 8–16 and 16–8 h. No aspect of growth investigated was associated with photoperiodic phenomena, and growth occuring during the light break (8-7.5-1-7.5 h) was intermediate between that in 8–16 and 12–12 h. Three patterns of cell elongation were found in the four species in which (1) cell age, (2) cell age and position and (3) cell age, cell position and light period determined cell length at different axial cell positions. Elongation was followed within cells, along axes ofAntithamnion spirographidis for plants grown under different day lengths. Three regions of development were found along main axes: (1) an apical region in which basipetal expansion was greater than acropetal expansion. (2) a zone of stability with equal elongation in apical and basal growth region of cells, and (3) a basal region with greater acropetal expansion. With increasing daylength, the zone of stability was extended to greater ranges of cell length.