Insurance against reproductive failure in a semelparous plant: bulbil formation in Agave macroacantha flowering stalks

Bulbils are small aerial rosettes that occur on the flowering stalks of semelparous Agave plants and in related families, and that are capable of acting as clones of the parent plant. We hypothesized that bulbil formation was inversely related to fruiting success in the flowering stalk, and we explored this hypothesis in A. macroacantha, a species from South-Central Mexico. Forty randomly chosen plants were divided amongst three treatments: (a) elimination of all floral buds, (b) exclusion of pollinators, and (c) control. We also studied 22 plants in which the flowering stalk had been felled by goat grazing. Between September and November 1991 we kept a record of the numbers of bulbils and capsules produced in each flowering stalk. Significant (P<0.0001) differences between treatments were found in the proportion of plants hearing capsules and bearing bulbils. The control treatment had the highest proportion of plants producing capsules, treatment a had the highest proportion of individuals bearing bulbils, while treatment b showed an intermediate response. In the goat-grazed group, 45% of the plants failed to produce any propagative structure after the stalk was cut, and half of all plants produced bulbils on the remaining stump. A significant inverse relationship between the numbers of capsules and the numbers of bulbils per plant was found for the three randomly assigned treatments. Our results suggest that once the production of the flowering stalk has been triggered and the death of the rosette is irreversible, bulbils may act as an insurance mechanism that increases the probability of successful reproduction of the genet.     

Dormancy in Dioscorea: Sprouting promotion by inhibitors of protein synthesis in bulbils and rhizomes

The sprouting of immature and half-dormant mature bulbils andrhizomes in some species of Dioscorea was promoted by treatmentwith inhibitors of protein synthesis, although the sproutingof completely awake mature bulbils was inhibited. Gibberellicacid (GA₃) inhibition of the sprouting was relieved by cycloheximideand some base and amino acid analogues. Polyphenol oxidase activity in the bulbils decreased after incubationwith the protein-synthesis inhibitors. Possible involvement of protein synthesis in inducing dormancyof Dioscorea bulbils is discussed. (Received July 11, 1977; )     

Dormancy in bulbils of several herbaceous plants: Effects of photoperiod,light, temperature,oxygen and gibberellic acid

Effects of some environmental conditions (photoperiod, white and colored lights, temperature, partial oxygen pressure) and growth regulators (gibberellic acid, 2-chloroethyltrimethylammonium chloride) on induction and release of dormancy of the bulbils ofDioscorea batalas, Laportea bulbifera, Elatostema involucratum andSedum bulbiferum were investigated. Bulbils were formed under short-day conditions inLaportea andElatostema, under long-day conditions inSedum, and irrespective of photoperiods inDioscorea. In all species exceptSedum, immature bulbils required light, particularly blue or far red, for sprouting (photo-sprouting stage), and mature bulbils required a cold treatment (thermo-sprouting stage). The duration of photo-sprouting and thermo-sprouting stages and the degree of dependency on light or low temperature of sprouting differed from species to species. Sprouting of chilled mature bulbils of these species was promoted by light, especially by red or green light. Both immature and mature bulbils ofSedum sprouted under short-day conditions. Continuous irradiation with blue, far-red and green light markedly inhibited their sprouting. Oxygen at high concentration inhibited the sprouting of immature bulbils inDioscorea; in the other species it promoted sprouting regardless of the maturation of the bulbils. Applications of gibberellic acid caused the sprouting of bulbils the absence of light, chilling or photoperiodic treatment in all species exceptDioscorea, in which gibberellic acid inhibited sprouting. Polyphenol oxidase activity was very high in the homogenates ofDioscorea bulbils, and increased further when the bulbils had been treated with gibberellic acid. In the other species, little or no such activity was observed.     

Dormancy inDioscorea: Different temperature adaptation of seeds,bulbils and subterranean organs in relation to north-south distribution

Temperature dependencies of sprouting and germination were compared for subterranean perennating organs and seeds of ten closely related species of the genusDioscorea (Dioscoreaceae), a group of monocotyledonous summer perennials which are distributed from the tropics to the subarctic. The species used wereD. nipponica Makino,D. tokoro Makino,D. japonica Thunb.,D. tenuipes Franch. et Savat.,D. septemloba Thunb.,D. quinqueloba Thunb.,D. izuensis Akahori,D. bulbifera L. f.spontanea (Makino) Makino et Nemoto,D. pentaphylla. L. andD. alata L.; they are distributed from cold northern areas to warmer southern areas approximately in this order in and around Japan. Bulbil sprouting was also studied in those forming bulbils. Subterranean organs of the tropical species sprouted faster without any prior temperature treatment, whereas those of species from the more northern areas sprouted after prechilling. Northern species required longer, periods of prechilling for sprouting. On the other hand, with seeds or bulbils, the southern species required longer periods of prior temperature treatment for dormancy breaking. This difference in the length of dormant periods between seeds or bulbils and subterranean organs among the ten species may be related to their size and position of shedding; seeds or bulbils are small and are shed on the ground surface, whereas subterranean organs are large and are located below the surface. It is important to determine in other perennials whether the above relation between dormant features of seeds or bulbils and subterranean organs are common properties or not.     

Studies on the growth and development of Oxalis latifolia

The growth and development of Oxalis latifolia was assessed on outdoor-grown potted plants raised from bulbs. Three main phases of growth appeared to be correlated with major physiological developments in the aerial and subterranean organs. Establishment (weeks 1 – 5) was characterised by a slow gain in plant dry weight accompanied by high relative growth rate (RGR) values for individual plant organs. The second phase of growth (weeks 6 – 12) was reproductive, as stolons grew and initiated bulbils at the apices. Bulbils had the capacity to produce their own shoot system and ultimately some primary bulbils supported secondary bulbils. Dry matter accumulation was in the order: bulbils > peduncles > leaves > petioles > stolons = roots > parent bulb. The RGR of individual plant organs declined together with the net assimilation (NAR) rates and the leaf area ratio of the shoots during the reproductive phase and beyond into the senescence phase (week 13 onwards). The results are discussed in relation to the problem of achieving weed control via cultural and herbicide treatments.     

Morphology and molecular phylogeny ofTretopileus sphaerophorus, a synnematous hyphomycete with basidiomycetous affinities

Tretopileus sphaerophorus, a synnematous hyphomycete with basidiomycetous affinities was newly isolated from the decaying petiole and peduncle ofCocos nucifera collected in Depok, Indonesia. The species produced first a bulbil as a propagule on the top of a synnema. After the bulbil had fallen, the synnema proliferated about seven times to produce new bulbils, each time making conspicuous nodes at the upper part. By careful morphological observation, clamp connections were confirmed on the hyphae in the specimens and culture. In culture, each hyphal cell with or without a clamp was found to be dikaryotic by DAPI nuclear staining. Germination of the bulbils occurred first from projecting hyphal tips on their upper surface, which have been treated as germ pores. The inner structure of the bulbils, the hyaline mucus of the bulbils, and conidium-like hyphal fragments were also examined. Phylogenetically,T. sphaerophorus was inferred to be related to the Aphyllophorales based on the nuclear encoded small subunit (18S) rDNA using the homology search system (FASTA) and the neighbour-joining method. Part 10 in a series on the taxonomy of synnematous fungi.     

Aspects of sexual failure in the reproductive processes of a rare bulbiliferous plant,<Emphasis Type="Italic"> Titanotrichum oldhamii</Emphasis> (Gesneriaceae), in subtropical Asia

Titanotrichum oldhamii produces both flowers and asexual bulbils on its inflorescences. However, field observations and herbarium collections indicate that seed set is infrequent and that most reproduction is from vegetative bulbils. We have investigated the failure of sexual reproduction and identified four major causes: (1) in the wild, the seed:ovule ratio for open pollination was only 1.9%, in contrast to 10.1% for artificial cross-pollination, implying poor pollinator services. (2) The overall reproductive success was 5–10 times greater in glasshouse pollination treatments compared to field treatments. This suggests suboptimal environmental conditions for seed set in natural habitats, which may eliminate natural seed set completely. (3) Pollination experiments showed that outcrossed populations set significantly more seed and had higher germination rates than intra-populational and self crosses, in both field and glasshouse experiments. T. oldhamii thus appears to benefit from wide outcrossing. Pollen transfer between populations, however, seems to be infrequent because of the rarity and scattered distribution of Titanotrichum populations. (4) Flowers near the apex of the inflorescence are less likely to set seed, especially late in the season when inflorescences convert to bulbil production. In these late flowers, pollen tubes showed poor guidance as they approach the micropyle of the ovules. Even under optimal glasshouse conditions, the average outcrossing success was only 0.229. Almost one-half of the ovules remained undeveloped and 13.5% of ovules aborted after pollination, indicating a strong shift of resource allocation toward vegetative bulbils and rhizome development. Efficient reproduction from asexual bulbils may thus have released Titanotrichum from strong selection for efficient sexual reproduction. However, occasional seed set observed in the wild may be very important for maintaining some genetic diversity in populations, and promoting overall fitness.     

A Landscape Perspective of the Stream Corridor Invasion and Habitat Characteristics of an Exotic (Dioscorea oppositifolia) in a Pristine Watershed in Illinois

The spatial distribution of exotics across riparian landscapes is not uniform, and research elaborating the environmental constraints and dispersal behavior that underlie these patterns of distribution is warranted. This study examined the spatial distribution, growth patterns, and habitat constraints of populations of the invasive Dioscorea oppositifolia in a forested stream corridor of a tributary of Drury Creek in Giant City State Park, IL. The distribution of D. oppositifolia was determined at the watershed scale mainly by floodplain structure and connectivity. Populations of D. oppositifolia were confined to the floodplain, with overbank flooding from the stream. Dioscorea oppositifolia probably originates in disturbed areas upstream of natural corridors, and subsequently, the species disperses downstream into pristine canyons or ravines via bulbils dispersing in the water. In Giant City State Park, populations of D. oppositifolia were distributed on the floodplain across broad gradients of soil texture, light, slope, and potential radiation. The study also examined the longevity of bulbils in various micro-environments to illuminate strategies for the management of the species in invaded watersheds. After 1 year, the highest percentages of bulbils were viable under leaves, and much lower percentages were viable over leaves, in soil, and in the creek (76.0±6.8, 21.2±9.6, 21.6±3.6, and 5.2±5.2%), respectively. This study suggests that management procedures that reduce leaf litter on the forest floor (e.g., prescribed burning) could reduce the number of bulbils of D. oppositifolia stored in the watershed.     

Physiological activity in persistent bulbils of Agave vilmoriniana (Agavaceae)

Plants of Agave vilmoriniana are unique members of a reproductively diverse genus since they produce prolific numbers of both bulbils and viable seeds. Bulbils were observed to remain attached to the flowering stalk into a second summer, instead of abscising in the year they developed. Hence, a number of hypotheses were tested to establish the physiological basis for their longevity and evaluate their influence upon the plant's reproductive biology. Bulbils produced following anthesis were abundant (>2000 bulbils), large (0.44 g dry mass per individual), and their combined leaf surface area (one side) could exceed 3 m². Thus, the current-year inflorescence may act as a photosynthate source. Second-year, attached bulbils are healthy-looking, even after leaves of the parent plant have senesced, since maternal roots continue to absorb rain water. Rehydrated 2nd-yr bulbils resume Crassulacean acid metabolism, sometimes demonstrating net daytime tissue acidification. Water conservation by bulbils is very efficient, exhibiting very low cuticular conductances (0.016-0.0035 mm/s). Bulbils detach over all climatic seasons after having formed >3 and >18 root primordia in their 1st and 2nd year, respectively. About one-third of bulbils are capable of forming roots after a 1-yr storage at 35°C, when their water contents averaged 37%. Irrigated bulbils produce water-absorbing roots within 2 d, with de novo chlorophyll synthesis and nocturnal acidification commencing within 4 d. Two bulbils were observed to root and become established in shaded microhabitats during this 2-yr study of 29 plants.     

Promotive and inhibitory effects of uniconazole and prohexadione on the sprouting of bulbils of Chinese yam,Dioscorea opposita

Gibberellin A₁ (GA₁), GA₃ and GA₄ inhibited the sprouting of nondormant bulbils of Chinese yam, Dioscorea opposita, where the effectiveness of the GAs was as follows: GA₄>GA₁+GA₃. Uniconazole and prohexadione, plant growth retardants, promoted the sprouting of half-dormant bulbils. By contrast, these retardants inhibited the sprouting of nondormant bulbils. Gibberellin A₃ (GA₃) and A₄ (GA₄) which were applied to the stems of the sprouted bulbils, promoted stem elongation, but GAs applied to the bulbous parts inhibited this process. The effectiveness of the GAs on stem elongation was as follows: GA₃+GA₄ for the promotion and GA₄ > GA₃ for the inhibition. Uniconazole applied to the stem inhibited the stem elongation of the sprouted bulbils. These results suggest the possible involvement of endogenous GAs in the induction and maintenance of bulbil dormancy of D. opposita, as well as in the bulbil sprouting and subsequent stem elongation.     

Response to competition of bulbous geophyte Allium oleraceum differing in ploidy level

Experimental studies that explore the possible causes of ploidy distributions and niche differentiation are rare. Increased competitive ability may be advantageous for survival in dense vegetation and may strongly affect local and regional abundances of cytotypes and potentially contribute to invasion success. We compared survival, growth and reproduction of plants originating from bulbils of three cytotypes (2n = 4x, 5x, 6x) of Allium oleraceum growing with and without a competitor (Arrhenatherum elatius). There was a strong negative effect of competition but no effect of ploidy or ploidy × competition on survivorship, height and total dry mass of A. oleraceum, i.e. no support for different competitive abilities of the ploidy levels. However, slightly different responses of populations to competition treatments within all cytotypes suggest differentiation within cytotypes. Under competition, plant survivorship was low, surviving plants were small, had low dry mass and produced neither sexual nor asexual propagules. Without competition, plant survivorship was high, and cytotypes differed in three traits after 2 year's growth: dry mass of flowers, number of flowers and ratio of the dry mass of sexual to asexual propagules all decreased with increasing ploidy level. We additionally tested tetra‐ and pentaploids as to whether plants originating from different types of propagule (bulbils, seeds) differ in survivorship, growth and reproduction when growing with and without a competitor. Plants originating from bulbils had higher survivorship, were more robust, flowered earlier and produced more propagules when compared to plants originating from seeds and grown without competition. Under competition, differences in performance between plants originating from seeds and bulbils mostly disappeared, with higher survivorship only for plants originating from bulbils.     

Identification of Endogenous Gibberellins and Abscisic Acid from Dormant Bulbils of Dioscorea japonica (Japanese Yam)

Four gibberelJins (GAs), GA₁₉, GA₂₀, GA₂₄, and GA₅₃, were identified by gas chromatography-mass spectrometry (GC-MS) from the dormant bulbils of Dioscorea japonica Thunb. ex Murray (Japanese yam), suggesting that two biosynthetic pathways of GAs, early 13- and non-13-hydroxylation pathways, are operating in the bulbils. Abscisic acid was also identified by GC-MS.     

Minor differences with big consequences: Reproductive patterns in the genus Gagea (Liliaceae)

Reproductive patterns in ten species of Gagea Salisb. were compared by counts and measurements of bulbs, bulbils and flowers in large cohorts including all life stages. Two types of bulbils were found: taxa with “type I bulbils” start to develop a single to several bulbils as soon as the replacement bulb has reached a certain diameter and then continue to form them indefinitely throughout the life of the plant. “Type II bulbils” are only temporarily produced in immature, non-flowering plants of some species, but not in fully grown, flowering individuals, a phenomenon termed “reproductive switch”. Patterns of bulbil formation are species-specific: G. davlianidzeae, G. nigra, G. peduncularis, G. pratensis, and G. spathacea produce only type I bulbils; G. angelae, G. fedschenkoana and G. lutea develop only type II bulbils. Both bulbil types occur simultaneously in G. fragifera and G. villosa. The quantitative investigations demonstrate the existence of species-specific thresholds for the development of bulbils as well as flowers. Compared to the adult volume of the replacement bulb (where 90% of all plants flower), both types of bulbils have usually low thresholds: 0–5% (type I, all but one species) and 3–13% (type II). Inflorescences develop if plants attain between 38 and 60% of the adult bulb volume. Minor changes in patterns of bulbil formation and thresholds for their development may ensure survival of highly sterile taxa (e.g. G. spathacea, G. fragifera). This, in turn, can facilitate speciation in the genus driven by both hybridization and polyploidization.     

A Bryophyte flora of Cornwall. II. Sphagna

Abstract

The taxonomy of Anomobryum julaceum and allied species with axillary bulbils in Europe and Asia is reviewed. A. concinnatum is regarded as a distinct species, occurring in W. and C. Europe, SW., N. and C. Asia and N. America. A. bavaricum has often been confused with A. concinnatum, but differs in its more numerous, small, reddish bulbils and in leaf shape; it is known only from the European Alps. The Asian A. nitidum also has numerous small reddish bulbils but it differs from A. bavaricum in leaf shape and bulbil form. A lectotype is designated for A. nitidum, of which A. gemmigerum and other nominal taxa are regarded as synonyms. Information is presented on geographical ranges and habitats of the four valid species.     

Dormancy in Laportea bulbils: experiments with buds cultured in vitro

Dormancy in bulbils of Laportea bulbifera was investigated usingbuds cultured in vitro. Buds excised from immature bulbils couldsprout in light, but scarcely in the dark. Regardless of lightconditions, buds isolated from mature bulbils failed to sproutwhile those from chilled mature bulbils could. GA and BA stimulatedthe sprouting of buds excised from dormant mature bulbils. Incontrast, ABA and morphactin inhibited the sprouting of budsexcised from nondormant chilled mature bulbils. These resultssuggest that, in dormant Laportea bulbils, the buds themselvesare in a dormant state which may be controlled by a balanceof activity between growth-inhibiting and -promoting substances. (Received October 29, 1976; )     

Effect of alar and CCC on induction and germination of bulbils in <Emphasis Type="Italic">Curculigo orchioides</Emphasis> grown in shake flask cultures

Bulbil formation in Curculigo orchioides is followed by asynchronous germination. The effect of alar and CCC incorporated in Murashige and Skoog (MS) medium has been studied on bulbil induction from leaf explants and subsequent germination of bulbils. MS medium contained 1 mg/l BA and 0.1 mg/l morphactin for bulbil induction while germination medium contained 1 mg/l gibberrelic acid and both the media contained alar or CCC (0.5–5.0 mg/l). Growth retardants markedly reduces the bulbil formation, yield and fresh weight of bulbils. Incorporation of retardants resulted in 60% germination inhibition, thereby prolonging the storage conditions.     

滴水珠珠芽发育过程研究

珠芽是滴水珠(Pinellia cordata)的营养繁殖结构,为揭示滴水珠珠芽发育过程中的特征,该研究以野外采集材料进行盆栽观察试验,通过形态观察法和石蜡切片方法,探索滴水珠叶片和叶柄处珠芽发育过程中的形态学和解剖学结构特征。结果表明:滴水珠珠芽发育过程在形态上分为叶柄无现象时期、早期珠芽结构分化和珠芽膨大成熟时期三个时期;相应的解剖学发育时期为珠芽原基启动期和形成期、原基分化期和膨大成熟期四个阶段。叶片和叶柄珠芽起始细胞均由叶柄腹面表皮下层薄壁细胞组织脱分化形成,起始细胞不断分裂形成细胞团最后发育成珠芽原基;在原基分化期的特征是生长点形成,最终分化形成芽原基和鳞片叶;膨大成熟期的特征是珠芽结构不断生长,生长点侧生分化鳞片叶增多。叶片和叶柄处珠芽成熟脱落时期为灰色椭圆形结构,测得平均直径分别为(2.79±0.40)mm和(2.69±0.36)mm,外部有鳞片包裹,内部含大量营养物质。在环境适宜条件下,珠芽遇土萌发,萌发率达75%。这表明滴水珠珠芽与同属植物半夏珠芽发育过程差别不大,都是无性克隆营养繁殖体结构。     

Loss of short-day requirement for dormancy breaking of bulbils from gibberellic acid-treated plants in Sedum bulbiferum

Sedum bulbiferum forms bulbils at superterranean nodes under long-day conditions, and the detached bulbils sprout after exposure to short-days [1]. When gibberellic acid was applied to the mother plant at the start of the long-day induction period, the number of bulbils formed increased slightly and these bulbils sprouted on incubation in the dark but not under short days of continuous light. However, when gibberellic acid was applied directly to detached bulbils during incubation, the short-day requirement for sprouting was conserved. Gibberellic acid application to the mother plants enhanced sprouting ability of detached bulbils when incubated under illumination with blue, green or far-red light. However, presence of gibberellic acid during bulbil exposure to light did not induced marked enhancement in sprouting under blue, green of far-red light. Thus, gibberellic acid application to the mother plant modified light and photoperiodic requirements for the sprouting of detached bulbils of S. bulbiferum.     

Causes of variation in sexual and asexual reproduction in diploid and triploid populations of <Emphasis Type="Italic">Allium scorodoprasum</Emphasis>

Allium scorodoprasum is a bulbous geophyte that reproduces by seeds, aerial bulbils, and subterranean daughter bulbs. This research represents the analysis of intraspecific variations in sexual and asexual reproduction in relation to ploidy levels and the environmental conditions from 11 diploid and 24 triploid populations in Lithuania. The results showed that the seed production was extremely rare and occurred only in diploid populations. The numbers of flowers and aerial bulbils per plant as well as mean mass of one bulbil were the characteristics in which differences between ploidy levels were the biggest. The diploids on average produced more (61.9), but smaller (mean mass = 29.6 mg) bulbils and more flowers (16.7), while triploids produced less (41.7), but larger (mean mass = 45.9 mg) bulbils and fewer (5.2) flowers. The study revealed that the main determinant of intracytotypic variation in quantity of both sexual and asexual structures was light availability.