Relations between submarginal and marginal sori in ferns

In contrast to the generally favoured hypothesis that the marginal position of sporangia and sori is the primitive condition, Hagemann (e.g. 1984a) holds the view that the superficial position of reproductive organs is primitive. Marginal reproductive organs correlate with a distinct succession of a vegetative and a reproductive developmental phase in the sporophyll (phase differentiation in leaf development). Since phase differentiation means an increase in complexity, marginal reproductive organs are considered advanced. In order to examine the relations between submarginal and marginal sori in the light of Hagemann's hypothesis, various soral forms of ferns were analysed using the methodological framework of construction morphology sensu Weber (1958). The sorus is regarded as a reproductive region of the leaf surface that comprises not only the sporangia but also the receptacle as an essential component. The growth pattern of the receptacle, the developmental succession of the sporangia, and the formation of the indusia lead to the recognition of several sorus types.In the present paper the basipetal marginal sorus is presented and defined as the combination of (a) a ridge-, cushion-, or rod-shaped receptacle, (b) a basipetal sequence of sporangia initiation, and (c) the presence of a lower and an upper indusium or of a cup-shaped indusium. As the first examples of this type, the sori of selected Hypolepidaceae and Dennstaedtiaceae are treated in detail. Regarding the phylogenetic relationships of the present families, the following conclusions can be drawn: (1) Owing to the meristem fusion of the indusia the sorus ofDennstaedtia possesses a new quality as compared to that of the Hypolepidaceae. Therefore, the Hypolepidaceae and the Dennstaedtiaceae should not be combined in a single family, as suggested by some authors. (2) The marginal position of the sori, and, in addition, the highly complex soral construction of the Dennstaedtiaceae, are not in support of the primitiveness of the two families, as sometimes assumed. (3) A sorus construction as found inDennstaedtia glauca can be gradually derived from a less elaborate one, such as that found, e.g., inPteridium. The continuous increase in the perfection of the soral construction (Pteridium Dennstaedtia) suggests a placement of the Dennstaedtiaceae after the Hypolepidaceae.     

Sorus development on median and marginal parts of the sporophyte of Laminaria japonica Areschoug (Phaeophyceae)

Sorus formation on median and marginal parts of Laminaria japoncia Areschoug was investigated by cultivating sporophytes from December 1996 to September 1997 in Southern Hokkaido, Japan. Simultaneously, discs (ca. 3 cm in diameter) that were cut off monthly from median and marginal parts of the cultivated sporophytes were incubated in our laboratory. All the cultivated sporophytes formed sori on the median parts from January to August, while all the median and marginal discs formed sori at 10 under 46 μmol photon m^-2s^-1 (12L:12D) in enriched medium. There was no difference in sorus area between median and marginal discs. These results indicated that the marginal parts are able to produce sori, although the greater part of them are actually washed away before forming sori in the sea. The marginal part is probably in a state of `resource limitation' on the reproduction because the resource outflowed from this part to the basal part for the growth of sporophytes. This revised version was published online in August 2006 with corrections to the Cover Date.     

Comparative photosynthetic studies of Ecklonia cava (Laminariales,Phaeophyta) bladelets with and without zoosporangial sori

Photosynthetic rates were compared between Ecklonia cava bladelets with and without zoosporangial sori sampled from the subtidal zone (about 5 m deep) in Nabeta Bay, Shimoda, Japan. Photosynthetic rates of bladelets were lower in the sorus portion than in the non-sorus portion on the basis of area, dry weight and chlorophyll a. Respiration rates were higher in the sorus portion than in the non-sorus portion on the basis of area and chlorophyll a, whereas they were almost the same on a dry weight basis. The differences were mainly due to a large difference in dry weight per unit bladelet area between the sorus and the non-sorus portion. Light compensation points were higher in the sorus portion than in the non-sorus portion.     

SPOROGENESIS UNDER ULTRAVIOLET RADIATION IN LAMINARIA DIGITATA (PHAEOPHYCEAE) REVEALS PROTECTION OF PHOTOSENSITIVE MEIOSPORES WITHIN SORAL TISSUE: PHYSIOLOGICAL AND ANATOMICAL EVIDENCE1

To study the effect of different radiation conditions on sporogenesis of Laminaria digitata (Huds.) J. V. Lamour., excised disks were induced to form sporangia under PAR (P), PAR + ultraviolet‐A (UVA) (PA), and PAR + UVA + ultraviolet‐B (UVB) (PAB) conditions in the laboratory. Vitality of meiospores, released from sori induced under different radiation conditions in the laboratory and from sori of wild sporophytes acclimated to in situ solar radiation in the presence and absence of ultraviolet radiation (UVR), was measured in terms of their germination capacity. Sorus induction in disks of laboratory‐grown sporophytes was not hampered under light supplemented with UVR, and sorus area was not significantly different among P, PA, and PAB. Vitality and germination rate of meiospores released from sori induced under different radiation treatments was comparable. Likewise, screening of UVR of the natural solar radiation did not promote higher germination rates of meiospores released from wild sporophytes. Germination rates were, however, higher in meiospores released from laboratory‐induced sori compared to sori of wild sporophytes. Higher DNA damage (formation of cyclobutane pyrimidine dimers, CPDs) was observed in laboratory‐grown nonsorus compared to sorus tissue, while CPDs were nondetectable in both sorus and nonsorus tissue of wild sporophytes. To explain the apparent protection of developing meiospores and the unexpected UV resistance of soral tissue, concurrent anatomical investigations of sporogenic tissue were performed. We observed the previously unreported existence of two types of sterile paraphysis cells. One type of paraphysis cells, the most frequent type, contained several red‐fluorescing plastids. The other type, less frequently occurring, was completely filled with substances emitting blue fluorescence under violet excitation, presumably brown algal phenolic compounds (phlorotannins). Cells of this type were irregularly scattered within the sorus and did not contain red‐fluorescing plastids. Meiospore‐containing sporangia were positioned embedded between both types of paraphysis cells. In vegetative tissue, blue autofluorescence was observed only in injured parts of the blade. Results of our study suggest that the sorus structure with phlorotannins localized in the specialized paraphysis cells may be able to screen harmful UVR and protect UV‐sensitive meiospores inside the sporangia.     

Phylogenetic relationships of ferns deduced from rbcL gene sequence

Part of the large subunit of the ribulose-1,5-bisphosphate carboxylase/oxygenase (rubisco) gene (rbcL) was sequenced from three fern species: Adiantum capillus-veneris, Botrypus strictus, and Osmunda cinnamomea var. fokiensis. This region included 1,333 base pairs, about 90% of the gene. Maximum likelihood analysis of the deduced amino acid sequences indicated that (1) Botrypus (Ophioglossaceae) clustered monophyletically with other ferns (Adiantum, Angiopteris, Osmunda); the closest relative to Botrypus among the three species was Osmunda, which did not support the hypothesis that the Ophioglossaceae are linked to the progymnosperm-seed plant lineage. (2) Eusporangiate ferns containing Botrypus (Ophioglossaceae) and Angiopteris (Marattiaceae) were a paraphyletic group. (3) Seed plants and the four fern species examined formed a monophyletic group, but ferns and bryophytes (liverwort) did not. Variations in rates of substitution for synonymous and nonsynonymous codons were found in fern lineages.Correspondence to: M. Hasebe     

Meiospores produced in sori of nonsporophyllous laminae of Macrocystis pyrifera (Laminariales,Phaeophyceae) may enhance reproductive output

Different lamina of Macrocystis pyrifera sporophytes (i.e., sporophylls, pneumatocyst‐bearing blades, and apical scimitars) in a wave‐sheltered site were found to be fertile. We quantified their sorus surface area, reproductive output (number of spores released) and the viability of released spores (germination rate). Sorus area was greatest on the sporophylls, with sporangia developing on >57% of the total area and smallest on the pneumatocyst‐bearing blades with 21% of the total area bearing sporangia. The apical scimitar released the greatest number of meiospores (cells · mL^?1 · cm^?2) and the sporophylls the least. Meiospores produced from all types of fertile laminae were equally viable. This reproductive plasticity may enhance reproductive output, and contribute to short and long‐distance spore dispersal and the cryptic gametophyte propagule bank for the next generation of sporophytes.     

BesitztAnemia Sori? Untersuchungen über die Entwicklung des Sporophylls vonAnemia phyllitidis (Schizaeaceae)

Observations of young fertile leaf primordia provide information about the development of the sporophyll ofAnemia phyllitidis Sw. The marginal meristem which surrounds the leaf primordium forms the pinna primordia, firstly the two “spore pinnae” by meristem fractionation. These are turned with their adaxial side towards the leaf apex and continue marginal meristem fractionations until products of the 5th order are formed.—In the sporophyll development two events are significant: (1) The fractionation products of the 2nd order reverse their direction of coiling. (2) From the marginal meristem of the fractionation products of the 5th order the sporangia arise in acropetal succession each originating from one initial cell.—Three observations—the fractionation products of the 2nd order being accessory outgrowths of the leaf margin, their reversed coiling direction, and the aggregation of the sporangia on the last segments—lead to the following concept of a sorus type: Each fractionation product of the 2nd order represents a marginal acropetal sorus with a branched receptaculum.     

Allocation of blade surface to reproduction in Laminaria longicruris of Long Island Sound (USA)

Sorus formation in the kelp Laminaria longicruris de la Pylaie (Phaeophyta) was quantified for a biennial population in Long Island Sound (Connecticut, USA), at the southern limit of its biogeographical range in the Western Atlantic Ocean. The allocation of blade surface to reproduction was measured monthly during two years, with additional samples taken at times of peak growth and reproduction. Sporophytes produced sori year-round, with the highest percentages of fertile plants (75 to 95 percent) occurring from October to December. The mean percentage of sorus area to blade ranged from 1 to 37 percent, remaining low and fairly constant during the spring and summer, but reaching much higher levels in the fall (when growth is minimal) and early winter. The comparatively low mean annual allocation of blade surface to reproduction (5 percent), was postulated to be due to temperature stress on these plants at the southern limit of distribution, but may instead be typical of the species. Such a small allocation of resources to reproduction nevertheless results in billions of spores per m² of substrate, making a major contribution of material potentially available to grazing food webs.     

Fertile organs and in situ spores of a new dipteridaceous fern Hausmannia sinensis from the Jurassic of northern China

As a representative fossil member of the dipteridaceous fern, genus Hausmannia was reported worldwide from the Mesozoic strata; however, little is known about the fertile structures, including sporangia and in situ spores, of this genus. In this study, a new species Hausmannia sinensis was identified from the Middle Jurassic of Nei Mongol (Inner Mongolia), northern China. The specimens are compressions and are well preserved with details of sporangia and in situ spores. The leaf laminae are broadly fan-shaped, with an almost entire margin. Primary and lateral veins dichotomously branch to form square or polygonal meshes. Each ultimate mesh bears one to two circular sori of 0.4 mm in diameter. Sori are exindusiate; each sorus contains three to six round to ovoid sporangia. The annulus is developed and oblique, with stomial region present in proximal position. Spores are trilete, circular to oval in shape. Both proximal and distal surfaces are covered with baculate to subverrucate sculptures. Spores range from 20 to 30 μm in diameter (average 28 μm), and are comparable to the dispersed genera Baculatisporites Thomas and Pflug and Apiculatisporis Potonié and Kremp. Hausmannia sinensis represents the first compression species of genus Hausmannia form Eurasia, which shows the combination of well-preserved sori, sporangia, annuli and in situ spore characters, and is therefore helpful for further understanding the diversity and evolution of the Dipteridaceae fern lineage through time.     

Cyathea cranhamii sp. nov. (Cyatheaceae), anatomically preserved tree fern sori from the Lower Cretaceous of Vancouver Island, British Columbia

Permineralized cyatheaceous sori occur among remains of conifers, fungi, and other plants in newly discovered calcareous concretions from Early Cretaceous (Barremian) marine sediments of Vancouver Island, British Columbia, Canada. Sori are superficially attached in two rows to narrow pinnules and display a globose sphaeropteroid indusium. Annulate sporangia with multicellular stalks diverge from a basal, vascularized receptacle. The nearly vertical uniseriate annulus is not interrupted by the stalk. The sporangia bear 64 trilete spores with perispore sculpturing that ranges from irregular granulate/echinate to prominent rodlets. These specimens, described as Cyathea cranhamii sp. nov., are the first anatomically preserved tree fern sori from the fossil record. They represent the most ancient evidence for fertile structures of the Cyatheaceae and demonstrate that essentially modern species of cyatheaceous tree ferns had evolved by the Early Cretaceous.     

Life history and reproductive potential of the agarophyte Gelidium robustum in California

The reproductive potential of the tetrasporangial phase of Gelidium robustum was studied for 16 months at two sites off Santa Barbara, California. In all samples tetrasporangial thalli were always more abundant than gametangial ones. Tetratrasporangial sori were present throughout the duration of the study but relative fecundity was highest [300–400 sori g^–1 (w. wt)] in spring/summer samples of consecutive years, as a result of increasing numbers both of tetrasporangial branchlets per plant and of sori per branchlet. On the other hand, laboratory experiments showed that tetraspore release per sorus was highest (150–250 spores sorus^–1 d^–1) in winter. Inferring from these field and laboratory data plants released up to ± 34 000 tetraspores g^–1 (w. wt) d^–1 in the spring/summer of the second study year. Tetraspore germination, under defined culture conditions, also showed a marked seasonality increasing sharply from less than 10% in winter up to almost 60% in spring/summer, thus coinciding with the period of maximal spore output per plant. These results suggest that although relatively high numbers of tetraspores may be released by G. robustum plants all year round these might not always have the potential to germinate and recruit.     

Development of the zoosporangia ofSynchytrium endobioticum,the causal agent of potato wart disease

Summary An ultrastructural study of zoosporangium development ofSynchytrium, endobioticum (Schilb.) Perc. is presented. Emphasis is placed on the location of the parasitic fungal thallus in the potato host cell, on the specific location of organelles in relation to the developing zoosporangial wall, and on the host cell reaction to the fungal infection. The cytoplasmic organization of the individual sporangia after division of the zoosporangium into a sorus of sporangia is characterized by numerous similarly sized nuclei, well developed dictyosomes, and the presence of many lipid bodies of variable size. Cytoplasmic microtubules are observed to flare out from the functional kinetosome both before and after zoospore cleavage.The ultrastructural details of zoosporangium development are used to revaluate the life cycle ofS. endobioticum as described from light microscopic observations made early in the century (Curtis 1921;Köhler 1923, 1932;Percival 1910).     

Isolated, distal blade discs of the brown alga Laminaria digitata form sorus, but not discs, near to the meristematic transition zone

Blade discs of vegetative thalli of Laminaria digitata (Huds.) Lamour. from Helgoland (North Sea) cut at 5–15 cm distance from the blade/stipe transition, formed sorus in the laboratory after 7–12weeks, 5 months earlier than whole fronds in the field. Sorus formation occurred in a broad range of daylength regimes or temperatures, at 8–16 h light pe rday and 6–12 ^°C. No sorus was developed during three months by meristematic blade discs cut from the lowermost 3 cm portion of the blade, nor from whole thalli cultured in parallel to isolated blade discs. These findings point to the possible existence of sporulation inhibitors produced by the laminarian meristem. This revised version was published online in August 2006 with corrections to the Cover Date.     

THE DEVELOPMENT OF THE SPOROCARP OF MARSILEA VESTITA

The tissues of the sporocarp of Marsilea vestita undergo profound changes during development. Early in development, the cells of the peripheral tissues, epidermis, hypodermis and layers of the transitional zone between the hypodermis and more internal tissues contain prominent vacuolar bodies. As development proceeds, these vacuolar bodies disappear. Prominent amyloplasts are found only in the guard cells and in the cells of the transitional zone. Later in development the cells of the hypodermis divide periclinally forming two layers which differentiate as macrosclereids. The cells of the outermost layer of the transitional zone differentiate as osteosclereids. Internally, the cells of the sorophore accumulate large amounts of mucilage in the central vacuoles. The peripheral cytoplasm ultimately degenerates leaving just hygroscopic mucilage. The mucilage carbohydrate contains the sugars, rhamnose and arabinose. In the young sorus, only the spore mother cells and the cells of the indusium contain amyloplasts. By the time of meiosis, there is a massive accumulation of starch in the receptacle, stalk and jacket but not in the tapetum of the sporangia. Late in development, the starch disappears and the mega- and microspores become coated with carbohydrate.     

Quantitative comparison of foliage development amongDryopteris monticola,D. tokyoensis and a putative hybrid,D. kominatoensis in Northern Japan

Developmental leaf architecture was quantitatively described in terms of measurements of various parameters on leaf blade from different size of sporophytes inDryopteris monticola, D. tokyoensis and a putative hybrid,D. kominatoensis in the natural site of Hokkaido, to compare the ontogenetic differentiation in foliage structure among allied ferns. The morphological stage of leaf and sporophyte was tentatively quantified by the number of midrib branches of the leaf (NV, number of veins), which exhibited a significant correlation to the leaf-shape complexity from a circle (DI=marginal length/2×(3.14×square)^1/2) of leaf blade. D. kominatoensis showed intermediate values between others in following characters; DI increase, maximum NV (also blade length), maximum number of costa branches of pinnae (NVMP), number of costa branches of the lowest pinna (NVLP), difference between NVMP and NVLP (NVMP-NVLP), during heteroblastic leaf development. A larger number of leaves per sporophyte was found inD. kominatoensis than in others. The fertility rate (%) and initiation of fertility (IF) in the relative developmental stage (RDS) ofD. kominatoensis shifted to that ofD. tokyoensis, while the order of pinnae with NVMP shifted to that ofD. monticola. Even in the intermediate characters inD. kominatoensis, slight shifts in characters to those of putative parents were found during heteroblastic leaf development. Contribution No. 3145 from the Institute of Low Temperature Science, Hokkaido University.     

Studies on the rust,Maravalia cryptostegiae,a potential biological control agent of rubber-vine weed,Cryptostegia grandiflora (Asclepiadaceae: Periplocoideae), in Australia,III: Host range

Seventy-one plant species, which were selected using a centrifugal phylogenetic method, were screened againstMaravalia cryptostegiae, an endemic rust of rubber-vine (Cryptostegia grandiflora) in Madagascar. All species and variants of the genusCryptostegia proved to be highly susceptible to the rust under optimum greenhouse conditions. Two other species belonging to the same subfamily Periplocoideae,Gonocrypta grevei from Madagascar andCryptolepis grayi from Australia, developed fertile sori but could still be classified as partially resistant, based on the results of a leaf clearing and staining technique. Fifteen assessment categories and 10 susceptibility ratings were designated by analysing both macro- and microsymptoms. Five resistance mechanisms were identified in the non-hosts and these are described and illustrated. Resistance reactions ranged from complete suppression of spore germination on the plant cuticle to restriction of sorus development due to the collapse of rust haustoria. The significance of host range studies are discussed in relation to the use of rust fungi as classical biological control agents of weeds, and specifically the release ofM. cryptostegiae in Australia.     

紫萁孢子囊发育中多糖和脂滴的细胞化学观察

采用光镜、透射电镜和细胞化学技术,对紫萁孢子囊发育过程中孢壁的超微结构和孢子囊内多糖和脂滴的分布及其动态变化进行研究,以探讨紫萁孢子囊发育过程中多糖和脂滴的代谢特征,为蕨类孢子发生的研究提供基础资料。结果表明:(1)紫萁孢子囊由1层囊壁细胞、2层绒毡层和产孢组织构成。(2)紫萁孢子壁由发达而分2层的外壁(外壁内层和外壁外层)和薄的不连续的周壁构成,由外壁形成棒状纹饰的轮廓;孢子外壁内层由多糖类物质构成,外壁外层和周壁均含有脂类物质。(3)在紫萁孢原细胞中观察到少量脂滴;随着紫萁孢壁的形成,囊壁细胞中淀粉粒的大小逐渐变小、数目先增加后减少,它们转运到内层绒毡层原生质团并转化为孢粉素前体物质,再穿过原生质团内膜表面进入囊腔,成为孢粉素团块或以小球形式填加到孢子表面形成孢壁。(4)紫萁孢子囊将多糖类营养物质转化为脂类,以脂滴的形式储藏在孢子中。     

Taxonomical studies on ovariicolous ustomycetes on caryophyllaceae I.Ustilago jehudana andUstilago moenchiae-manticae

A study of the type specimen ofUstilago jehudana resulted in the correction of the diagnosis. The sori are localized in the host gynoecium but not in the anthers. Morphological characters of the sori and ustospores of the later describedU. moenchiae-manticae are identical with these ofU. jehudana. Ustilago moenchiae-manticae is reduced here to a synonym ofU. jehudana. The smut is reported as new to Bulgaria on a new host, viz.,Moenchia erecta. A new combination,Bauhinus jehudanus, is proposed.     

Sorophores of<Emphasis Type="Italic">Lygodium</Emphasis> Sw. (Schizaeaceae) from the Late Cretaceous of New Jersey

We report here on a series of specimens of charcoalified sorophores with characteristics of the extant fern genusLygodium (Schizaeaceae) collected from sediments of the Raritan Formation (Late Cretaceous). Each elongate lobed fertile pinnule (sorophore) is flattened and bears alternately arranged sporangia on one surface. Each sporangium is covered by an indusium continuous with the margin of the lamina. Sporangia are oblong in shape, short stalked, and have an apical annulus formed by a single ring of radiating cells that dehisces longitudinally. The sporangial cap or distal face is formed by only one cell. All of these features are characteristic of the extant genusLygodium. Small numbers of trilete, psilate spores are found in the sporangia. Megafossils assignable toLygodium are known from the Upper Cretaceous of North America and Germany with worldwide distribution during the Tertiary. The newLygodium fossils are compared with others previously referred to the genus.