Photoperiodic regulation of receptacle induction in Sargassum horneri (Phaeophyceae) using clonal thalli

We developed a clonal culture of Sargassum horneri to investigate the effect of photoperiod on reproduction in this species. Regenerated vegetative thalli were obtained using lateral branches excised from a thallus grown from a single embryo under short‐day conditions (SD = 10:14 h light : dark cycle). Lateral branches excised from the SD‐regenerated thallus became vegetative thalli that remained in that phase as long as they were cultured under SD. When an excised lateral branch was cultured under long‐day conditions (LD = 14:10 h light : dark cycle), it began to enter the reproductive phase while still less than 50 mm long. Induction of the reproductive phase was accompanied by a distinctive morphological change – suppression of blade formation at the apical region of the branch; elongation of branches without blades was then followed by differentiation of receptacles bearing conceptacles on their surface. Apices of receptacles were able to interconvert between reproductive and vegetative phases, as blades resprouted upon transfer from LD to SD. The critical day length for induction of receptacle formation was between 13 and 14 h; receptacle formation was also induced under SD conditions with night breaks (NBs). These results strongly suggest that reproductive regulation of S. horneri is a photoperiodic long‐day response. NBs with blue and green light were effective for reproductive induction but not with red light. This suggests that blue‐ and/or green‐light photoreceptors are involved in the photoperiodic reproductive response of S. horneri.     

REPRODUCTION AND LIFE HISTORY OF THE RED ALGA GALAXAURA OBLONGATA (NEMALIALES,GALAXAURACEAE)1

Culture and morphological studies showed that Galaxaura oblongata (Ellis et Solander) Lamouroux has a triphasic life history with conspicuous gametophytes and small filamentous tetrasporophytes. Development of male and female reproductive structures is very similar and both begin with the enlargement of a terminal cell of a filament branch occupying a normal vegetative position within the apical pit of a thallus branch. In male thalli this modified branch forms a conceptacle in which spermatangia are produced. In female thalli, this modified branch forms a three-celled carpogonial branch consisting of a carpogonium, hypogynous cell and basal cell. Filament branches from the basal cell form a pericarp and the gonimoblast develops directly from the carpogonium. Carposporangia are produced in conceptacles which resemble the male conceptacles. About the time the first carposporangia are produced, the carpogonium, hypogynous cell and basal cell form a large fusion cell. Released carpospores germinate in a unipolar or bipolar manner and form small filamentous thalli. Under short day conditions, cruciate tetrasporangia are produced in small clusters. Tetraspores germinate similarly to carpospores and also form small filamentous thalli. Under low nutrient conditions, small cylindrical thalli develop on the filaments and these appear similar to gametophytes collected in nature.     

Organismic determinants and their effect on growth and regeneration in Gracilaria gracilis

The growth of Gracilaria gracilis (Stackhouse) Steentoft, Irvine et Farnham was examined by studying the effect of organismic determinants such as thallus length, position along the thallus and branching. Knowledge of these factors is essential in order to increase production from suspended seaweed rafts seeded with vegetative G. gracilis fragments. Seeding netlons with seaweed material freshly collected from subtidal populations provided up to 30% higher relative growth rates than seaweed maintained on the netlons for successive months. Initial seedstock length greatly affected growth rates and yields such that 30-cm thalli fragments resulted in growth rates 14% higher than for 10-cm fragments. This difference is suggested to be due to the higher contribution to overall biomass by growth of lateral branches. Comparisons of the growth of apical and basal fragments suggest that growth takes place over the entire length of the thallus, but that the apex contributes more to overall elongation than does the proximal part. The removal of apical meristems resulted in an enhanced branching frequency with production of four times as many branches as intact fragments. Evidence is also provided for extensive morphological differentiation following long periods of rapid growth. These thalli have very high frequency of branching, are hollow due to the disintegration of medullary cells and are considered to be completely senescent. These factors have implications for the successful cultivation of G. gracilis on commercial mariculture systems. This revised version was published online in July 2006 with corrections to the Cover Date.     

Lobe formation and division in the foliose lichen <Emphasis Type="Italic">xanthoparmelia conspersa</Emphasis>

New lobe development and lobe division was studied in the foliose lichen Xanthoparmelia conspersa (Ehrh. ex. Ach.) Hale. In thalli with either the centre or margin removed, the inside edge of the perimeter, the outer edge of the reproductive centre, and fragments derived from the thallus perimeter all regenerated growing points (‘lobe primordia’) within a year. Thalli possessing isidia had the greatest ability to regenerate growing points. In reproductive thalli, there was a positive correlation between the density of new growing points and thallus size. When fragments were cut from the perimeters of mature X. conspersa thalli and glued to pieces of slate, the ratio of growing points to mature lobes increased over 54 months. Lobes within a thallus exhibited different degrees of bifurcation. In some bifurcating lobes, the point of origin of the bifurcation advanced at the same rate as the lobe tips over 4 months but in most lobes, the bifurcation point either advanced less rapidly than the lobe tips or retreated from its original location. Removing adjacent lobes had no significant effect on the radial growth of a lobe over 4 months or on the location of the bifurcation point but it increased the number of growing points. These results suggest that for X. conspersa: 1) all portions of of thalli can regenerate growing points, 2) few growing points actually develop into mature lobes, 3) individual lobes within a thallus grow and divide differently, and 4) adjacent lobes inhibit the development of growing points on their neighbours.     

Phototropism of Thalli and Rhizoids Developed from the Thallus Segments of Bryopsis hypnoides Lamouroux

Newly regenerated thalli were used to study the phototropism of Bryopsis hypnoides Lamouroux under different qualities of light. Positive phototropism in the thalli and negative phototropiam In the rhizoida of B. hypnoides were investigated and analyzed in terms of bending. Both thaiii and rhlzoids developed from thallus segments exhibited typical tip growth, and their photoreceptive sites for phototroplam were also restricted to the apical hemisphere. The bending curvature of rhizoids and thalli were determined with unilateral lights at various wavelengths and different fluence rates after a fixed duration of Illumination. The trends of bending from the rhizoid and thallus were coincident, which showed that the action spectrum had a large range, from ultraviolet radiation （366.5 nm） to green light （524 nm）. Based on the bending curvatures, blue light had the highest efficiency, while the efficiency of longer wavelengths （〉500 nm） was significantly lower. External Ca^2＋ had no effect on the bending curvature of thalli and rhlzolda. Blue light （440 nm） induced thallus branching from rhizoids, while red light （650 nm） had no such effect. Fast-occurring chloroplast accumulation In the outermost cytoplasmic layer of the blue light （440 nm）-Irradiated region In the rhizoid was observed, from which protrusions （new thalli） arose after 4 h of the onset of illumination, and this action was thought to be driven by the dynamics of actin microfilamenta.     

浒苔(Enteromorpha prolifera)藻体发育的显微观察

通过显微镜观察以及石蜡切片法观察了浒苔属浒苔藻体的发育过程,阐述了浒苔从孢子释放到形成成熟藻体这一发育过程的显微特征。结果表明:浒苔孢子由成熟体细胞分化发育而成,每个成熟体细胞可形成10～30个孢子;孢子首先分裂为2细胞结构,2细胞具有明显的极性,基细胞发育成为假根,顶端细胞不断进行横分裂形成丝状体;丝状体顶端以下细胞通过纵分裂形成管状叶状体,顶端细胞始终保持横分裂;纵分裂分为切向分裂与径向分裂,切向分裂增加管状叶状体管周细胞数从而使管体增大,径向分裂形成管外壁突起细胞,最终发育为分枝;管状叶状体管腔内部有绒毛状的突起及糖蛋白成分的网架结构;当管状叶状体管周细胞达到30～50h,管腔内部突起消失,网架结构收缩拉紧,管壁细胞贴近形成片状叶状体;整个浒苔藻体始终保持极性发育。     

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.     

Competitive equivalence in a community of lichens on rock

Lichens and mosses cover 70–100% of the rock surface in a forested Appalachian boulderfield, and competition for space is intense. This paper examines overgrowth ability and its morphological correlates in four common species of foliose lichen on rocks. Overgrowth requires one lichen thallus to overtop another at the point where they meet. Therefore, I quantified margin height for a number of thalli of each of four lichen species. Two umbilicate species attached to the rock only at the thallus center showed a positive relationship between thallus size and margin height: large thalli often reached considerable heights above the rock surface, yet most also had points along their margin that were quite low and flat. Two other nonumbilicate species were characteristically flatter and showed no dependence of margin height on thallus diameter. Differences among species, among thalli of the same species, and among different points on a single thallus accounted for approximately equal amounts of variance in margin height. To determine the success of species in overtopping each other, I then recorded 639 instances of apparent overgrowth (overtopping of one thallus by another) on several rocks in the boulderfield. Of the nine pairs of species that met often enough to permit statistical analysis, only four pairs showed a consistent winner. Species in the remaining five pairs were competitively equivalent, neither winning significantly more than half the encounters although each individual encounter had a clear winner. Overgrowth rates measured from sequential photographs were highly variable, but many species pairs showed no substantial differences between growth rates over another thallus and growth rates over bare rock; only one species appeared to be affected by overgrowing other thalli. Overgrown thalli, as well as thalli experimentally shaded by gluing an overhanging rock chip above their margin for a year grew very slowly or not at all in the region of overgrowth, and the overgrown region of the thallus was often markedly discolored or disintegrating. Thus, foliose lichens compete strongly for space at the study site, yet because competitive success is at least partly based on a morphological character (margin height) that is inherently quite variable even in a single thallus, many pairs of species appear to be competitively equivalent.     

QUANTITATIVE VARIATION IN VULPINIC AND PINASTRIC ACIDS PRODUCED BY TUCKERMANNOPSIS PINASTRI (LICHEN-FORMING ASCOMYCOTINA,PARMELIACEAE)

Tuckermannopsis pinastri is a lichen species found commonly on rocks and tree branches in boreal and alpine habitats in the northern United States. Members of this species produce three yellow-pigmented phenolic compounds: usnic, pinastric, and vulpinic acids. The objective of our study was to quantify the variation in concentration of the latter two of these compounds in relation to substrate factors, chlorophyll content, and thallus size. Using high-performance liquid chromatography, we analyzed 120 thalli of T. pinastri collected randomly from a single large population located at Spruce Knob, West Virginia. Although individuals were sampled from both tree and rock substrates that differed markedly in light intensity, these environmental factors were not correlated with observed variations in vulpinic or pinastric acid concentrations. Instead, compound concentrations were correlated most closely with thallus size, with small rather than large thalli having the highest concentrations of the two compounds. Small thalli did not have higher concentrations of chlorophylls than large thalli, however, which suggests that the rate of production of secondary compounds by the fungus in T. pinastri is independent of algal biomass. Inasmuch as lichen secondary compounds serve a defensive role against microorganisms and herbivores, our results suggest that small, juvenile thalli are better defended than more mature thalli.     

Unique distribution of epiphytic Neosiphonia harveyi (Rhodomelaceae,Rhodophyta) along sargassacean hosts

Although epiphytism is generally heaviest on older portions of the host thallus and both variation and abundance of epiphytes decrease with proximity to meristematic apices, Neosiphonia harveyi (Bailey) Kim, Choi, Guiry and Saunders was frequently found on upper parts of sargassacean hosts. This study compared density, thallus length and fertility of epiphytic N. harveyi among different regions of Sargassum patens C. Agardh thalli to reveal a unique distribution pattern of this epiphyte. The majority of epiphytic N. harveyi was observed on the upper part of host thalli and was seldom seen near the basal part. Over a 3‐month period, mean epiphyte density was greater on host apices while thallus length and fertility were greater on subapical portions of host thalli, suggesting the colonization by this epiphyte near the apical part of the host. Distribution patterns of this epiphyte were similar among S. patens growing at different depths. Apical portions of S. patens appear to be more suitable substrata for N. harveyi settlement and colonization compared with other portions of the host thallus, regardless of depth.     

Intrathalline and size-dependent patterns of activity in Lasallia pustulata and their possible consequences for competitive interactions

1. In dense populations of the saxicolous lichen Lasallia pustulata the margins of adjacent thalli overlap each other in intraspecific competition for space and light.
2. In situ non-destructive monitoring of hydration-dependent potential photosynthetic activity by modulated fluorescence systems in different parts of the thallus shows that the activity is structured by a centre-to-margin gradient, with the centre of the thallus remaining active for substantially longer periods than the margins when the thalli dry up after being activated by wetting. The pattern reflects the water status of different parts of the thallus; the margins which are thin and exposed dry up first.
3. The activity pattern within individual lichen thalli suggests that marginal overlapping between neighbours may have a less detrimental effect on the shadowed individuals than expected from a pure consideration of the amount of area shadowed. Because the centre of the lichen thallus is active for longer periods, shadowing of this region may possibly be more harmful per area unit than an overlap at the less active margins.
4. Larger thalli are active for substantially longer periods than small ones. Even the margins of larger thalli tend to be active for a longer period than the centre of small thalli.     

Growth of foliose lichens: a review

This review considers various aspects of the growth of foliose lichens including early growth and development, variation in radial growth rate (RaGR) of different species, growth to maturity, lobe growth variation, senescence and fragmentation, growth models, the influence of environmental variables, and the maintenance of thallus symmetry. The data suggest that a foliose lichen thallus is essentially a ‘colony’ in which the individual lobes exhibit a considerable degree of autonomy in their growth processes. During development, recognisable juvenile thalli are usually formed by 15 months to 4 years while most mature thalli exhibit RaGR between 1 and 5 mm yr⁻¹. RaGR within a species is highly variable. The growth rate-size curve of a foliose lichen thallus may result from growth processes that take place at the tips of individual lobes together with size-related changes in the intensity of competition for space between the marginal lobes. Radial growth and growth in mass is influenced by climatic and microclimatic factors and also by substratum factors such as rock and bark texture, chemistry, and nutrient enrichment. Possible future research topics include: (1) measuring fast growing foliose species through life, (2) the three dimensional changes that occur during lobe growth, (3) the cellular changes that occur during regeneration, growth, and division of lobes, and (4) the distribution and allocation of the major lichen carbohydrates within lobes.     

Developmental morphology of the thalloid Hydrobryum japonicum (Podostemaceae)

We describe the unique development and branching of lobed thalli in Hydrobryum japonicum. Lobe formation begins with meristem initiation at random sites near the thallus margin fringed by protective tissues. As the protective tissues are successively peeled off particularly in the growing new lobes, the lobes become naked and then become fringed again by new protective tissues that develop from the marginal part of the new meristems. Subsequently the meristems become less active and are differentiated into parenchymatous ground tissue at maturity. The random pattern of meristem formation during the sporadic development gives rise to a nonorderly branching pattern of the thalli. Some other lobes (～10%) are regenerated from injured parts of the thalli. The vegetative shoots arise endogenously near the thallus margin and are enclosed by the nonvascular strand nets. The rudimentary shoot apices remain embedded in the thalli. The thalli, though remarkably different from typical roots of other angiosperms, might be extremely transformed roots.     

CHANGES IN WATER ECONOMY IN RELATION TO ANATOMICAL AND MORPHOLOGICAL CHARACTERISTICS DURING THALLUS DEVELOPMENT INPARMELIA ACETABULUM

The influence on uptake and water loss of the structural changes experienced by Parmelia acetabulum during thallus development were investigated. Small specimens were characterized by flat lobes and a thin thallus and cortex. Large specimens appeared strongly rugose, imbricate and irregularly folded, and had a significantly thicker cortex and medulla than small thalli. Maximum water storage capacity did not differ between large and small thalli, although water retention was much higher in large thalli, presumably due to the interaction of structural characteristics and a higher boundary layer resistance. This translated into a longer duration of the period of thallus hydration in large thalli compared to small thalli. Incubation of thalli in water-vapour-saturated atmospheres induced full recovery of photosynthetic electron transport to the values before desiccation in small thalli but only induced a partial recovery in large thalli. The close correlation between photosynthetic electron transport and net photosynthesis during desiccation found in this species suggested that carbon-fixation activity could be regained to a larger extent by incubation of thalli in water vapour in small compared to large thalli. The higher ability for water vapour uptake of small thalli might allow them to efficiently use small amounts of intermittently available water or periods of high relative humidity. The significance of this differential ability to utilize water is discussed with regard to the known ecological preferences of the species.     

Branch development in Lupinus angustifolius L.#II. Relationship with endogenous ABA, IAA and cytokinins in axillary and main stem buds

The concentrations of indole-3-acetic acid (IAA), cytokinins (CK) and abscisic acid (ABA) were measured in buds of different regions (main stem and lateral branches) of Lupinus angustifolius L. (cv. Merrit) and at different stages in the development of branches. In lupin, branching patterns are the result of discrete regions of axillary branches (upper, middle and basal) which elongate at much different rates. Early in development only the main shoot elongates, followed usually by basal branch growth and then rapid upper branch growth. Branches in the middle of the main stem grow only weakly or fail to develop. Levels of IAA were generally high in the apical buds of slowly growing branches and low in buds from strongly growing branches, whereas CK levels showed the opposite relationship. CK:IAA ratio showed a closer relationship with the rate of growth of a particular branch better than the levels of either CK or IAA alone. During early stages of growth ABA concentration did not follow the rate of branch growth. However, later in development, where growth did not closely match the ratio of CK:IAA, ABA level showed a strong negative relationship with growth. A significant decrease in ABA was associated with continued strong growth of the main stem apex following a decline in CK:IAA ratio. Overall, the best relationship between the level of growth factors in apical buds and branching pattern in lupin was the ratio of CK:IAA, implying that high CK:IAA at a given bud would promote growth. ABA level appeared to play a secondary role, as a growth inhibitor.     

Some observations on polarity and regeneration in Enteromorpha

The regeneration of two types of Enteromorpha has been investigated. Small tubular sections cut from the thallus develop rhizoids along the basal cut edge and papillae from the apical cut edge. The capacity for regeneration is greatest in segments from the middle and base of the thallus and least in apical sections. Regeneration in various liquid culture media at different light intensities and temperatures and on solid agar media has been tested. The addition of growth substances and extracts from Enteromorpha thalli always stimulate regeneration but in no way alter the polarity. The results are compared with previous conflicting accounts of regeneration of Enteromorpha.     

Growth and Dormancy in Lunularia cruciata (L.) Dum.: I. RESUMPTION OF GROWTH AND ITS CONTINUATION

Lunularia cruciata may become dormant at three stages in itslife history: mature thallus, gemma, and spore. The resumptionof growth and its continuation in various conditions have beenstudied in thalli and gemmae. Air-dry, mature thalli of theIsrael strain, planted on a suitable medium, produce adventitiousbranches ventrally from the region immediately posterior tothe existing meristem, which itself fails to resume growth.When dormant gemmae are taken from the gemma-cup, however, theexisting apical cells and meristems simply recommence growing,so that the new growth is continuous with the old. Except inthe case of mature thalli aroused from dormancy, apical dominanceis pronounced, and branching occurs only by bifurcation. Thisdominance can be broken by applying sucrose solution to thegrowing tips—possibly a plasmolytic effect. The growth in area of freshly planted gemmae accelerates forabout 25 days before its relative rate slows to any great extent.During the first half of this period, growth is due exclusivelyto the expansion of existing cells, but subsequently cell numbersincrease rapidly from the 8–10,000 present in the dormantgemma. Cell numbers were estimated by using a modified macerationtechnique, in which chelation followed prolonged fixation. Lunularia grows successfully at quite low light intensities.Of the mineral nutrient deficienccs investigated, lack of Plimits growth most severely, although N-deficiency also restrictsit to a very low level. Gemma-cup production appears to be unaffected by light intensity,at least within levels permitting growth. There is, however,a large temperature effect, cup production decreasing markedlyabove 12° C.     

The significance of thallus size for the water economy of the cyanobacterial old-forest lichen Degelia plumbea

Rosette-formed, circular thalli of Degelia plumbea were studied in the laboratory. Regardless of thallus size, the optimal quantum yield of photosystem II (F _V/F _M) remained at a high, constant level during a drying cycle starting with fully hydrated thalli until the thallus water content reached about 200%. Net photosynthesis reached a maximum level at this hydration level. Thereafter, both F _V/F _M and net photosynthesis fell rapidly to zero at a water content of somewhat less than 100%. There was a highly significant, positive relationship between thallus size and the water-holding capacity, as well as a strong, negative correlation between size and water loss per thallus area. Consequently, an increase in thallus size from 1 to 36 cm² lead to a tenfold prolongation of the photosynthetically active period during a drying cycle at a low radiation regime. The improved water-holding capacity in larger thalli is mainly a result of a thicker hypothallus. The fast desiccation of small thalli suggests that the regeneration of D. plumbea could be severely hampered by nearby logging that raises the evaporative demand by increasing radiation loads and wind exposure at remaining lichen sites. Received: 12 December 1997 / Accepted: 20 March 1998     

DISPERSAL PATTERNS OF TERRICOLOUS LICHENS BY THALLUS FRAGMENTS

Dispersal patterns from seven terricolous lichen species, with a high capacity for asexual reproduction by fragmentation (Cetraria muricata, Cladonia species), were studied in differing vegetation types in north-eastern Germany. Marked lichen thalli were fragmented by trampling. After 15 days the spread of thallus fragments was monitored. Whereas most of the fragments that were dispersed by wind remained within a 20-cm radius from the source, the maximal dispersal distance was 57 cm in a dry sand grassland and 68 cm in an open pioneer pine forest. Dispersal was negligible in a closed old-growth pine forest. Several fragmented lichen cushions were disturbed and removed by animals, and led to a maximal dispersal distance of 9·70 m. These results suggest that: (a) thallus fragments provide good short-distance dispersal in open vegetation, but are inefficient for long-distance dispersal, and (b) wind and animals are important factors for the dispersal of thallus fragments. For restoration management of man-made substrata, artificial introduction of lichen thalli is proposed.     

鹧鸪菜生殖器官发育的研究

鹧鸪菜[Caloglossa leprieurii(Mont)J.Ag.]细胞含多核。在生殖季节,孢子体、雌、雄配子体在外形上有明显的差异:四分孢子体大而平展,分枝的上部肉眼可见有红色小斑点,即为四分孢子囊群。雌配子体顶部略蜷曲,分枝密,囊果呈球状,多位于分枝上部分枝点的腹面。雄配子体个体较小,枝顶蜷曲,颜色淡,特别生有精子囊的部分更淡。每种藻体的营养细胞的纹孔连结非常发达,每个细胞都具有4-5条,多者达7-8条。果胞枝由四个细胞组成,一般由围轴细胞分裂形成,果胞受精后,由支持细胞分裂形成辅助细胞,一个生育小枝上,可以连续形成5-7个甚至10个以上幼果胞系,但多数仅有一个果胞受精发育成囊果。当一个果胞受精时,其他果胞系就停止发育。精于囊位于分枝上部中肋的两侧,背腹面均有分布。一个营养细胞沿着叶面平行方向平周分裂成三个细胞,在两个外侧细胞各形成3-4个精子囊母细胞,每个精子囊母细胞斜壁分裂产生2-4个精子囊,放散后,余下的精子囊母细胞仍留在中间盘状细胞上。