Climate, size and flowering history determine flowering pattern of an orchid

The flowering pattern of plant species, including orchid species, may fluctuate irregularly. Several explanations are given in the literature to explain that pattern, including: costs associated with reproduction, herbivory effects, intrinsically triggered unpredictable variation of the system, and external conditions (i.e. weather). The influence of age is discussed, but is difficult to determine because relevant long-term field observations are generally absent in the literature. The influence of age, size, reproductive effort and climatic conditions on flowering variability of Himantoglossum hircinum are examined using data collected in a long-term project (1976–2001) in Germany. PCA and multiple regression analysis were used to analyse variability in flowering pattern over the years as a function of size and weather variability. We studied future size after flowering to quantify costs of reproduction. Flowering probability was strongly determined by plant size, while there was no significant influence of age class on flowering probability of the population. Costs associated with reproduction resulted in a decrease in plant size, causing reduced flowering probability of the plants in the following year. The weather explained about 50% of the yearly variation in the proportion of large plants and thus had an indirect, strong influence on the flowering percentage. We conclude that variability in flowering is caused mainly by the variability of weather conditions in the previous and current year, whereby reproductive effort causes further variability in flowering at the individual and, consequently, the population levels.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 511–526.     

A spatially explicit analysis of seedling recruitment in the terrestrial orchid Orchis purpurea

Seed dispersal and the subsequent recruitment of new individuals into a population are important processes affecting the population dynamics, genetic diversity and spatial genetic structure of plant populations. Spatial patterns of seedling recruitment were investigated in two populations of the terrestrial orchid Orchis purpurea using both univariate and bivariate point pattern analysis, parentage analysis and seed germination experiments. Both adults and recruits showed a clustered spatial distribution with cluster radii of c. 4-5 m. The parentage analysis resulted in offspring-dispersal distances that were slightly larger than distances obtained from the point pattern analyses. The suitability of microsites for germination differed among sites, with strong constraints in one site and almost no constraints in the other. These results provide a clear and coherent picture of recruitment patterns in a tuberous, perennial orchid. Seed dispersal is limited to a few metres from the mother plant, whereas the availability of suitable germination conditions may vary strongly from one site to the next. Because of a time lag of 3-4 yr between seed dispersal and actual recruitment, and irregular flowering and fruiting patterns of adult plants, interpretation of recruitment patterns using point patterns analyses ideally should take into account the demographic properties of orchid populations.     

Flowering frequency in a small population of Gymnadenia conopsea-a five year study

This investigation evaluates the usage of genetic markers, microsatellites, to distinguish and re-identify individual plants in a population of the orchid Gymnadenia conopsea. The study also illustrates the problem in estimating the size of a population from single year sampling as individuals can rest underground or occur in vegetative states; information extremely important for the understanding of population dynamics as well as providing information for conservation management. The total population size was, based on information from microsatellite loci, estimated to 84 individuals and vastly larger than the annual number of flowering plants (mean 31.4 individuals/year). Flowering frequency varied from 24–49%, 53 individuals flowered once, five individuals were flowering four years and a single individual was flowering five years. A common pattern was one or two flowering periods followed by a non-flowering period. The observed number of alleles and the observed and expected heterozygosity varied among loci, although allele frequencies and genotype frequencies did not vary significantly among years.     

Spatial genetic structure in populations of the terrestrial orchid Cephalanthera longibracteata (Orchidaceae)

Orchid seeds are unusual for being the smallest among flowering plants. These dust-like seeds are wind-borne and, thus, would seem to have the potential for long-distance dispersal (a common perception); this perception has led to a prediction of near-random spatial genetic structure within orchid populations. Mathematical models (e.g., simple ballistic model) for wind-dispersed seeds and wind-tunnel experiments, in contrast, indicate that most seeds of orchids should fall close to the maternal plant (<6 m), supporting a prediction of significant fine-scale genetic structure within populations. In reality we do not know much about seed dispersion in orchids. To determine which of these two predictions is more appropriate, Wright's F statistics and spatial autocorrelation analysis were used to examine the genetic structure within two adult populations of the terrestrial orchid Cephalanthera longibracteata (Orchidaceae) in southern Korea. In results comparable to those of other self-compatible, mixed-mating plant species, C. longibracteata populations exhibited low levels of genetic diversity (mean H(e) = 0.036) and a significant excess of homozygosity (mean F(IS) = 0.330), consistent with substantial inbreeding via selfing and/or mating among close relatives in a spatially structured population. Spatial autocorrelation analysis revealed significant positive genetic correlations among plants located <10 m, with relatedness at <3 m comparable to that expected for half sibs and first cousins. This genetic structure supports the prediction that the majority of seed dispersal occurs over distances of less than 10 m and is responsible for generating substantial overlap in seed shadows within C. longibracteata populations.     

Orchid biology: from Linnaeus via Darwin to the 21st century

Orchidaceae are the largest family of flowering plants, with at least 24 000 species, and perhaps better than any other family of flowering plants, orchids represent the extreme specializations that are possible. As a result, they have long fascinated luminaries of the botanical world including Linnaeus and Darwin, but the size of the family has historically been an impediment to their study. Specifically, the lack of detailed information about relationships within the family made it difficult to formulate explicit evolutionary hypotheses for such a large group, but the advent of molecular systematics has revolutionized our understanding of the orchids. Their complex life histories make orchids particularly vulnerable to environmental change, and as result many are now threatened with extinction. In this Special Issue we present a series of 20 papers on orchid biology ranging from phylogenetics, floral evolutionary development, taxonomy, mycorrhizal associations, pollination biology, population genetics and conservation.     

Pollination efficiency and reproductive patterns in relation to local plant density,population size,and floral display in the rewarding Listera ovata (Orchidaceae)

Pollination efficiency and reproductive success vary strongly among populations of most animal‐pollinated plant species, depending on their size and local density, whereas individual plants within populations experience varying levels of reproductive output as a result of differences in floral display. Although most orchid species have been shown to be severely pollination limited, few studies have investigated the impact of the above‐mentioned factors on pollination success and reproduction, especially in rewarding species. In this study, the impact of population size, local density of flowering plants, and floral display on the rates of pollinia export and fruit production was investigated in 13 natural populations of the rewarding terrestrial orchid Listera ovata. In addition, an emasculation experiment was set up to examine how floral display and local density of flowering plants affected the relative importance of cross‐ vs. geitonogamous pollination in determining fruit set. In the studied populations, pollination efficiency, pollen removal, and fruit set increased with increasing population size until a threshold value of 30–40 flowering plants was reached, above which pollination efficiency and reproductive output decreased again. On average, plants with large floral displays showed higher proportional pollinia removal and fruit set compared with smaller plants. Fruit production was also significantly and positively related to local plant density, whereas emasculation did not affect the relationship between local plant density and fruit set, suggesting that geitonogamous pollination did not affect the outcome of female function. The results of this study are discussed in the light of the flowering mechanism of the species and its generalized pollination system. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 713–721.     

Seedling establishment in an epiphytic orchid: an experimental study of seed limitation

Will increased fruit and seed production in a severely pollination-limited orchid stimulate population growth? We tested whether safe sites for germination and seedling establishment are limiting for the twig epiphyte, Tolumnia variegata, by manipulating fruit set and monitoring subsequent seedling establishment for two seasons (1991–1992, 1992–1993). In the Cambalache Forest Reserve of Puerto Rico, we established 36 plots along a transect. Each plot consisted of nine trees. A center tree was designated as the site for attaching Tolumnia and manipulating fruit set. The other eight potential host trees were 1–3 and 3–5 m from the center tree in each of the cardinal directions. A 1-m length of stem 1 m from the ground was monitored for recruits on each of the nine trees of 24 fruit-enhanced plots and 12 controls (23 and 13, respectively for the 1992–1993 season). Fruit enhancement plots were divided among two treatments: one-fruit and five-fruit additions for the 1st year and one to five and more than five fruits for the 2nd year. Availability of suitable host species was not limiting. T. variegata showed little specificity for host tree species, good host trees and shrubs were common, and there was no evidence that the orchid had a preference for small branches, despite possessing the entire suite of characteristics thought to respresent “obligate” twig epiphytes. Fruit enhancement increased seed rain and seedling establishment consistently in only the high-fruit treatment plots. Most recruitment occurred near fruiting plants. Over the 2-year period, mortality was 18% for adults and 85.5% for the 1991–1992 cohort of recruits. Net recruitment was positive for both the treatment (average = 1.74) and control plots (average = 0.67). Seedling establishment at our study site was not microsite-limited. If selection for increased pollinator attraction occurs, then an increase in seed output should result in population growth.     

Symbiotic seed germination of Habenaria macroceratitis (Orchidaceae), a rare Florida terrestrial orchid

The rapid loss of native orchid habitat throughout ecologically important areas (e.g., Florida) has prompted researchers to develop appropriate plans for the propagation and reintroduction of many native orchid species. Ideally, symbiotic orchid seed germination methods are utilized in the production of orchid seedlings to be used in plant reintroduction programs. In the current study we (1) describe an efficient symbiotic seed germination protocol to germinate seeds of the rare sub-tropical terrestrial orchid Habenaria macroceratitis; (2) discuss the in vitro fungal specificity demonstrated by this species; and (3) describe the effects of three photoperiods (0/24 h, 16/8 h, 24/0 h L/D) on in vitro symbiotic seed germination of H. macroceratitis. Six fungal mycobionts were isolated from both vegetative and flowering plants of H. macroceratitis from two geographically distinct sites. Symbiotic seed germination percent was highest (65.7%) and protocorm development was most advanced (Stage 2) when seeds were cultured with fungal mycobiont Hmac-310. Seeds of H. macroceratitis demonstrated a degree of specificity toward fungal mycobionts isolated from plants originating from the same site where seed was collected. Continual darkness (0/24 h L/D) inhibited initial seed germination (Stage 1; 17.1%), but stimulated subsequent protocorm development (Stage 2; 53.5%). These findings will aid in developing an efficient symbiotic seed germination protocol for the conservation of this rare Florida terrestrial orchid, and may prove useful in the conservation of other sub-tropical terrestrial orchid species.     

Systemic endopolyploidy in <Emphasis Type="Italic">Spathoglottis plicata</Emphasis> (Orchidaceae) development

Background  

Endopolyploidy is developmentally regulated. Presence of endopolyploidy as a result of endoreduplication has been characterized in insects, mammals and plants. The family Orchidaceae is the largest among the flowering plants. Many of the members of the orchid family are commercially micropropagated. Very little has been done to characterize the ploidy variation in different tissues of the orchid plants during development.     

The unique pollination ofLeporella fimbriata (Orchidaceae): Pollination by pseudocopulating male ants (Myrmecia urens,Formicidae)

Leporella fimbriata is a self compatible orchid of southern Australia. It is dependant across its range on unique pollination by sexually attracted male winged antsMyrmecia urens, which pseudocopulate with the flower. Typical pollination sequences began with an initial circling then zig-zag flight to the flower. Vectors usually alighted on the inflorescence stem and quickly crawled to the flower where they adopted a copulatory position sideways along the wide labellum, pseudocopulatory probing immediately followed. In this position pollen carried on the thorax was deposited on the stigma. Departure from the labellum usually resulted in pollinium removal. Pollinator movements were restricted and the distribution leptokurtic with a mean of 3.141 ± 4.59 m. Pollination was widespread but variable from site to site and season to season with a maximum of 70% of all flowers being pollinated. Pollinator limitation is indicated. Traits essential for this pollination interaction include the coincidence of orchid and ant geographic distributions and the coincidence of flowering with the flight period of the ant. The production of pheromonelike substances and the distinctive floral morphology are also essential for attraction and manipulation of male ants. The ant mating system which the orchid can exploit is also important.     

Effects of floral display and plant abundance on fruit production of Ryncholaelia glauca (Orchidaceae)

Flowering plant density can increase number of visits and fruit set in multi-flowering plants, however this aspect has not been studied on few flower species. We studied the effects of individual floral display and plant density on the fruit production of the epiphytic, moth-pollinated orchid, Ryncholaelia glauca, in an oak forest of Chavarrillo, Veracruz, Mexico. Species is non-autogamous, and produced one flower per flowering shoot each flowering season. We hypothesized that orchids with more flowering shoots and those on trees with clumps of conspecific should develop more fruits than isolated ones. R. glauca population flowers synchronouly, and individual flowers last up to 18 days, with flowers closing rapidly after pollination. Individuals produced few flowers per year, although some plants developed flowers in both seasons and fewer of them developed fruits both years. There was no relationship between flower number per orchid, or per host tree, with the number of fruits developed per plant. Host trees with flowering and fruiting orchids were randomly dispersed and the pattern of distribution of flowering and fruiting plants was not related. Apparently, pollinators visit the flowers randomly, with no evidence of density dependence. The fruit set of R. glauca was as low as fruit set of multi-flowered orchids moth pollinated, suggesting that fruit set on moth pollinated orchids could be independent of the number of flowers displayed.     

Appearance and disappearance of a weedy orchid,Epipactis helleborine

Mark-recapture techniques applied to individual orchid plants and to their populations can further our understanding of the factors affecting establishment and survival. Studies need not be limited only to species at risk of extinction. Common or even introduced species may be tracked to determine which variables most impact behaviour and to develop effective conservation management techniques. Only long tracking periods are likely to intercept critical stochastic events. A 20-year study (1985–2004) of the introduced orchid,Epipactis helleborine, in Gatineau Park, Québec, Canada, has provided insight into the behaviour of this weedy species. We have marked 1591 plants over this period, 62% of which emerged only once. High late-season rainfall had a positive effect on emergence patterns whereas spring drought had a negative impact. The annual emergent population declined from a maximum of 315 plants in 1987 to 23 plants in 2001 following a severe drought in 1991. Twenty-seven plants, all but one of which grow in the vicinity of a large tree, re-emerged after prolonged absences of 7 to 18 years. A group of 15 plants that have shown complex patterns of re-appearance over the last 10 years also encircle this tree while 8 other plants with similar patterns of re-appearance but have not been seen since 1994 were distributed elsewhere in the site. We postulate that the ecological effect of large trees may be critical to the persistence of mycorrhizal associates of both the trees and the orchids and hence contribute to the extended hypogeal survival of the orchid.     

Does community‐level floral abundance affect the pollination success of a rewardless orchid,Calanthe reflexa Maxim.?

Decreases in pollinator abundance may particularly constrain plants that lack floral rewards, since they are poor competitors for pollinators in the plant community. Here, we documented the pollination ecology of a rewardless orchid, Calanthe reflexa Maxim., and examined effects of forest understory degradation by deer browsing on pollination success of the species in the light of a change in the abundance of neighboring flowering plants in 2010 and 2011. Bombus species were the only pollinators at each site and the flowering phenology of C. reflexa did not overlap with that of other rewarding plants. Pollinator visit rates (assessed by time‐lapse photography), and pollinia removal rate were higher in the undegraded understory site than the degraded site in both years, while the fruit set ratio did not differ between the sites in 2011. Coverage by neighboring flowering plants was extremely low in the degraded site. Our results suggest that, although its flowering phenology and consequently lower interspecific competition of C. reflexa with rewarding plants for attracting bumblebees, neighboring flowering plants may play an important role for maintaining the visitation frequency of bumblebees of C. reflexa and contribute to its pollination success.     

Mast fruiting in a hawkmoth-pollinated orchid Habenaria glaucifolia: an 8-year survey

Aims The hypothesis of predator satiation has been proposed to explain mast fruiting in various flowering plants. It considers that the simultaneous production of large numbers of seeds by a plant population reduces the risk of seed predation for each individual. Orchids produce huge numbers of seeds per fruit and rarely experience seed predation. It remains unclear which factors may affect fluctuating fruit production in orchids, which generally suffer a widespread pollen limitation. To explore the temporal pattern of fruiting and potential factors related to fluctuation in fruit production, we investigated reproductive success of a long-spurred orchid (Habenaria glaucifolia) in an alpine meadow with thousands of individuals over 8 years.Methods To estimate reproductive success, pollinator observation was conducted by day and at night, and pollinia removal and receipt were recorded in the field population for 8 years. To examine whether fruit set and seed set are pollen limited, we conducted supplementary pollination experiments and compared fruit set, seed set and pollinia movement of open-pollinated flowers from 2011 to 2013. We measured lengths of spurs and pollinator proboscises, and nectar volume and concentration, to identify potential pollinators.Important findings Hawkmoths were seen to be effective pollinators for H. glaucifolia in 3 years, whereas in the remaining 5 years no pollinators were observed, and consequently pollinia were rarely transferred. Numerous pollinia movements were observed in 2012, 2013 and 2014 (pollinia removal: 48, 59 and 85%; pollinia receipt 51, 70 and 80%), and correspondingly fruit set was significantly higher in 2012 and 2013 (59 and 46%) than in 2011 (25%). It was fruit set, rather than seed set, that was pollen limited in this orchid in the 3 years, in that supplementary pollination increased fruit set but did not increase seed set per fruit compared to natural. Three species of hawkmoths had proboscis lengths that matched the spur length of H. glaucifolia. Fruit set in this long spurred orchid depends on the activity of long-tongued hawkmoths, resulting in significant temporal variation in fruit production. Mast fruiting in this alpine orchid could be attributed to a 'sit and wait' strategy, awaiting an abundance of effective pollinators.     

Aspects of the population biology of Liparis loeselii (L.) Rich. var. ovata Ridd. ex Godfery (Orchidaceae) in the dune slacks of South Wales, UK

Demographic techniques have been employed to study populations of the rare orchid Liparis loeselii var. ovata across a range of dune slack habitats at a South Wales site. The study has shown that very large numbers of shoots (>300) may occur within a single 1 times 1 m quadrat and that the vegetative production of side shoots is common. Regular population surveys over time have shown that individuals represented by a single shoot can live for 8 years. Subsequent analyses of the floristic composition of the study quadrats has revealed the presence of subtle variations in species composition and vegetation structure in relation to age and hydrology, although all of the plots still fall within the SD14 Salix repens-Campylium stellatum dune slack community defined by the British National Vegetation Classification. When attributes of population flux are analysed in relation to floristic composition, one significant trend to emerge is that the appearance of new genets is most prevalent in successionally young slacks. As the natural formation of new slacks by sand blow is now a rare event in the South Wales dune field, the management of Liparis loeselii var. ovata is currentiy aimed at attempting to ensure the persistence of suitable communities through biomass reduction techniques.     

Research on orchid biology and biotechnology

Orchidaceae constitute one of the largest families of angiosperms. They are one of the most ecological and evolutionary significant plants and have successfully colonized almost every habitat on earth. Because of the significance of plant biology, market needs and the current level of breeding technologies, basic research into orchid biology and the application of biotechnology in the orchid industry are continually endearing scientists to orchids in Taiwan. In this introductory review, we give an overview of the research activities in orchid biology and biotechnology, including the status of genomics, transformation technology, flowering regulation, molecular regulatory mechanisms of floral development, scent production and color presentation. This information will provide a broad scope for study of orchid biology and serve as a starting point for uncovering the mysteries of orchid evolution.     

兰花花发育的分子生物学研究进展

兰科植物是开花植物中最大的家族之一,其花高度进化,具有花瓣状的萼片,特化的唇瓣和雌雄蕊合生的蕊柱,是单子叶植物花发育生物学研究的理想材料。近年来有关兰花花发育基因调控的研究已取得了一些进展,本文从兰花开花转换和兰花花器官的形成两方面综述了近年来国内外关于兰花花发育分子机理方面的研究进展,主要介绍了文心兰、蝴蝶兰和石斛兰的花发育相关基因,并推测了兰花花被的进化发育过程,认为兰花的DEFICIENS(DEF)类基因在早期经过两轮复制,形成了四类DEF基因,从而促进了花萼与花瓣的分离、侧瓣与唇瓣的分离。该文最后对今后兰花花发育研究的发展方向进行了展望。     

Population dynamics and life-history of Coeloglossum viride (L.) Hartm.: an endangered orchid species in The Netherlands

Coeloglossum viride(L.) Hartm. has a worldwide distribution in the Northern hemisphere where it can be found in a broad range of habitats; it is not, at present, threatened with extinction. However, in several European countries there has been a dramatic decline in sites occupied by this orchid. In The Netherlands only 2 populations remain of 55 extant before 1950. Knowledge of its ecology and life history are necessary to preserve this species at sites not directly threatened by habitat destruction. The results of a 7 year demographic study of a population ofC. viridein The Netherlands are presented. Annual recruitment and mortality vary considerably from year to year, whereas the flux in individual plants is always high. The half-life of the cohorts varies from 1.0 to 2.4 years and is on averagec. 1.5 years. This indicates thatC. virideis a short-lived terrestrial orchid species. The number of flowers per inflorescence varies from 2 to 26 per spike. The mean flowering percentage in the population is rather high (50%), while the fruit production was low in the year this was recorded (1994),viz. c. 10%. The average number of seeds per capsule wasc. 2300. Plants emerging above ground for the first time are already able to produce an inflorescence, and the average number of flowers equals those of plants initially passing a vegetative life phase for one or more years. Dormancy is a rare phenomenon in this species and the absence above ground when it does occur does not last longer than one growing season. Vegetative multiplication plays a minor role in population dynamics. The studied population is likely to be in a healthy state: there is a high recruitment almost every year resulting in a vigorous age structure. Management of the site, with mowing in July following seed dispersal and sheep grazing in winter is beneficial. The absence of intentional fertilization is a condition for a suitable, low vegetational structure of the sward during the growing season of this species.     

Trading populations—can biodiversity offsets effectively compensate for population losses?

Biodiversity offsetting promotes the protection or restoration of biodiversity at one site to compensate for the loss of biodiversity due to development at another site. Thus populations of species at a development site may be extirpated in the belief that offsetting elsewhere will compensate for the loss of biodiversity. In this study we tested the replaceability of roadside populations of the orchid Diuris platichila threatened by development (populations 1–5, n = 50–541 plants) with a potential offset population occurring in nearby natural vegetation (population 6, n = 143 plants). We measured differences in habitat among the populations and associated differences in flowering and fruiting. We also measured genotypic diversity within and among the populations, and the capacity of soil from each population to promote the symbiotic germination of outcrossed seed from the two largest populations (populations 1–2). An evaluation of the performance of the relevant offset policy was also undertaken which was informed by these studies. Compared to the roadside populations, the potential offset site had limited flowering (except after fire) and was genotypically less diverse. Soil from the potential offset site, and populations 2, 3 and 5, supported significantly less seed germination than soil from population 1. Translocating individuals from the most genotypically diverse populations into the offset population could help to buffer against the loss of genetic diversity if offsetting was required; however, the limited reproduction and recruitment opportunities at the offset site could result in the eventual erosion of any initial increase in diversity. The offset policy failed to secure a suitable offset for a hypothesized loss of plants. More generally we conclude that offsetting approaches which do not assess genotypic diversity and recruitment capacity may fail in their objective of protecting species and that maintaining populations in safe sites may be required.     

Postpollination nectar reabsorption in the African epiphyte Aerangis verdickii (Orchidaceae)

Nectar reabsorption in flowering plants seems to be a rare phenomenon and to our knowledge it has not been reported previously in the Orchidaceae. In this study we present data that show statistically significant differences in nectar sugar concentrations before and after pollination. Virgin flowers of the African epiphytic orchid Aerangis verdickii showed mean sugar concentrations in the nectar of 14.4 and of 18.3% in 1994 and 1995, respectively (sucrose mass by mass equivalent). Sugar concentration in nectar from naturally pollinated flowers had a mean of 3.2 and of 8.7% for the same period. In an experiment in the wild 60 flowers were protected against nectar theft by the ants. Half of those flowers were hand-pollinated. Control (virgin) flowers had nectar with a mean value of 13.8%, while flowers measured 48 h after pollination had a mean sugar concentration of 4.3%. The mean volume of nectar based on measurements of 23 virgin flowers from nine different plants was 19 uL. We estimated an average energy value for the nectar produced of 684 J per plant per season. At the study site, over 60% of the unmanipulated flowers were robbed of all their nectar by arboreal ants, Polyrachis spp. (Formicinae). The adaptive significance of nectar reabsorption for A. verdickii is probably a function of the environmental stresses to which it has been exposed and the relative costs of nectar production.