Flowering and Fruiting Phenology of Magnolia hypoleuca

Abstract The floral biology of Magnolia hypoleuca , a tall tree species widely distributed in Japan, was studied in a deciduous broad-leaved forest in Hokkaido near the northern limit of distribution of the species. The flower is large, protogynous and nectarless. Its petals close after the female stage and again after the male stage. The mean duration of flowering of an individual flower is 3–4 days, while the flowering period of an individual tree lasts up to 40 days. The main pollinators of the flowers were beetles. Success of pollination is assumed to be affected by weather conditions at anthesis. Average fruit sets (ratios of fruits to flowers) were 13–25%. Hand-pollination of 44 flowers produced a significantly higher fruit set (28 fruits) than control flowers (33/133). These results suggest that the fruit set is limited by pollination. The frequency of infertile ovules was higher in controls than in hand-pollinated fruits, indicating that the seed set was also limited by pollination. We propose a mimicry hypothesis in which the non-rewarding female-stage flower mimics the rewarding male-stage flower to explain the short life-span of an individual flower, the long flowering period of an individual tree, and the coexistence of flowers of various stages on a single tree.     

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.     

Reproductive phenology in three Genisteae (Fabaceae) shrub species of the W Mediterranean Region

We studied the flower duration, the phenology of flowering and fruiting, and flower, fruit, and seed production in three Genisteae shrub species of the W Mediterranean Region: Cytisus multiflorus, C. striatus , and Retama sphaerocarpa . Flower duration was negatively correlated with temperature, and in the case of C. striatus it was also influenced by pollination. In Cytisus multiflorus , which flowers during winter in the population studied, two floral morphs were recognized differentiated by flower size, phenological pattern, and production of the reproductive organs: morph LF (large flowers) and morph SF (small flowers). The former is earlier in phenology but its reproductive success is less than the second since it not only produces significantly fewer flowers/plant, but also a smaller crop of fruit and seeds. The flowering of the LF morph lasted some three months and of the SF morph 11 weeks. The winter-spring flowering C. striatus , with a four month duration of flowering time, is sympatric with C. multiflorus and their flowering periods overlap, but the former is more successful reproductively. Retama sphaerocarpa is clearly spring flowering, with an extremely short duration of flowering (c. 6.5 weeks), but, unlike the other two species which disperse their seeds in the same season in which they are produced, it staggers the dispersal of its diaspores over more than one year. Despite the great flower per plant production, especially in Retama sphaerocarpa , and of the number of fruit initiated, the final crop of fruit and seed/plant is low in the three species (C. multiflorus : fruit 8.40% and seeds 1.96%; C. striatus : fruit 13.09% and seeds 4.12%; Retama sphaerocarpa : fruit 5.65% and seeds 1.33%).     

WITHIN- AND AMONG-TREE VARIATION IN FLOWER AND FRUIT PRODUCTION IN TWO SPECIES OF CARYA (JUGLANDACEAE)

The patterns of variability in the production of staminate inflorescences, pistillate flowers, and fruits for Carya ovata (Mill.) K. Koch and C. tomentosa (Poir.) Nutt. at Hutcheson Memorial Forest (Somerset County, New Jersey) were examined over a four-year period. We sought to determine 1) the patterns of variability in flowering and fruiting (within-trees, among-trees, and between-years), 2) if variable flowering might account for observed nut-bearing patterns, and 3) what were the relative contributions of intrinsic (genetic) and/or extrinsic (environmental) factors in determining flower production and fruit set. In general, our fine-scale analysis of shoots within canopies did not reveal a distinct mast fruiting pattern. While the number of male and female flowers varied significantly (among trees and between years), fruit set was not markedly affected. Variability of flowering and fruiting among shoots within trees was low. Both flowering and fruiting were observed to have a strong genetic and/or microenvironmental basis; however, flowering appeared more plastic and thus more sensitive to yearly environmental vagaries than did fruiting. Favorable environmental conditions at the time of flower differentiation may result in increased flower production; but, this does not necessarily result in increased fruit set. Many pistillate flowers abscise around the time of pollination and fertilization, apparently adjusting fruit set to available shoot resources. Approximately 50% of the pistillate flowers of both species fail to set fruit. Overall, individual trees exhibit their own flowering and fruiting schedules, suggesting the need to account for this level of variability in future studies of mast fruiting.     

南方红豆杉响应不同传粉式样的结实表现

采用开花前雌株短枝套袋隔离传粉、花期人工授粉、风媒传粉 对照 及其传粉障碍等几种传粉式样 ,比较不同传粉 授粉 方式对南方红豆杉 Taxus chinensis var.mairei 结实的影响 ;在风媒传粉条件下 ,研究了南方红豆杉不同种植行向、树冠不同方位、上下内外不同层面的结实表现 .结果表明 :南方红豆杉存在孤雌生殖现象 ,平均单枝结实率为 2 8.7% ,与风媒传粉相比较 ,存在极显著差异 P<0 .0 1 ;人工授粉能显著提高结实率 P<0 .0 5 2 0 .7% ;2～ 3月份开花期 ,江汉平原盛行东北风 ,在风媒传粉条件下 ,东西向种植的南方红豆杉有更多的授粉机会 ,其结实情况极显著优于南北向种植的 P<0 .0 1 ;东西向种植的南方红豆杉树冠果实的空间分布 ,受传粉条件包括风向、冠层方位及距传粉源的距离、冠层内外上下层次等因素的影响 ,方位间的结实量按东南西北方位依次为 :2 30 .375粒、185 .6 2 5粒、12 8.813粒、10 5 .4 38粒 ,呈递减趋势 ,冠外层结实量普通高于冠内层 ,树冠东向下外层结实量最高 4 16 .75粒 ,北向下内层最低 2 2 .0 0粒 .     

Phenological variation in fruit characteristics in vertebrate-dispersed plants

Summary We investigated inter-specific variation in fruit characteristics — fruit size, seed number per fruit, seed weight, nutritional content, fruit persistence, and fruit synchronization — in relation to flowering and fruiting phenology in 34 species of fleshy fruited plants. Except for aspects of fruit synchrony and persistence, the results in general were inconsistent with previous suggestions about adaptive variation in phenologically related fruit traits. The main results were as follows: (1) Late flowering, late fruiting, lengthy development time from flower to fruit, and highly persistent fruits constitute a complex of correlated characteristics among the species. (2) Synchronization of fruiting within individuals increased from early ripening fruits to late ripening fruits. Fruiting synchrony was more pronounced in species with a small crop size than in species with a large fruit crop, whereas synchrony was not significantly related to flowering synchronization, nor to life form. (3) Nitrogen and carbohydrate content of fruit pulp did not vary in relation to phenology, whereas lipid content decreased from early to late ripening fruits. (4) No seasonal trends were found for variation in seed size or seed number per fruit. (5) Interactions with flowering phenology and developmental constraints are important in phenological fruiting patterns. Temporal variation in start of fruiting was partly (36%) explained by variation in flowering time. Seed weight variation explained 17% of variation in development time from flower to fruit. (6) Despite constraints from flowering and seed development, some adaptive adjustment in fruiting phenology is likely to be allowed for among the investigated species. Such an adaptive variation in fruiting phenology was suggested by intra-generic comparisons of Prunus and Vaccinium species.     

Temporal and spatial variation in flower and fruit production in a food-deceptive orchid: a five-year study

Deceptive orchids are generally characterized by low levels of fruit set; however, there may be substantial variations in fruit set between sites and years. Within a single population, individual plants may also differ greatly in their reproductive output as a result of differences in inflorescence size or local density. In this study, we determined flower and fruit production over 5 years in two populations of the food-deceptive orchid, Orchis purpurea . All plants were monitored annually for survival and flowering at each site to determine whether flowering and fruiting induced costs. The number of flowers per inflorescence varied considerably from year to year (min: 36.6, max: 49.5). Average fruit set was low (7%) and varied considerably among years and populations. A considerable proportion of plants also failed to set any fruit. However, the probability of producing at least one fruit was not affected by inflorescence size or local density. The number of fruits was significantly related to inflorescence size, but proportional fruit set was not. Local density also did not affect the number of fruits, nor proportional fruit set. There was also no evidence that plants with large inflorescence size or high fruiting success had a larger probability of remaining vegetative the year after flowering than plants with small inflorescence size or low fruiting success. Our results suggest that pollinator-mediated selective forces on inflorescence size through female reproductive success alone are weak, most likely because of the low overall level of visitation and the resulting uncertainty of pollination at the individual level. Our results further demonstrate that investigation of patterns of fruit set over several years is needed to better understand the variability in female reproductive success that is typical of most plant–pollinator interactions.     

Floral resources,pollinators and fruiting in a threatened tropical deciduous tree

Aims Nothapodytes nimmoniana (Family Icacinaceae) is a deciduous tree species distributed in Asia facing severe population decline. Wood chips from the tree are a source of camptothecin, a globally sought-after alkaloid with cancer-treating properties, and are harvested unsustainably in natural forests. We studied the pollination ecology of the species and asked if there are constraints in pollination and fruiting success in its natural populations. We also discuss the potential effects of wood extraction on pollinators and reproductive success in the population.Methods The study was carried out during the flowering season in two populations with varying exploitation levels, both located in the Protected Area Network in the Western Ghats of Karnataka State. We assessed floral resource availability and measured pollinator diversity and activity in the canopy from the perspective of the biology of the species. We quantified pollinator abundance and percent fruit set as a function of population density.Important findings Flowers belong to the fly pollination syndrome and are open to visits by generalists such as Apis dorsata and Trigona iridipennis. Fruiting success did not vary between exploited and unexploited sites, but there was a significant difference in pollinator assemblage. In a lean male flowering year, fruit set was low, suggesting pollen limitation in the population. No fruiting was recorded in the second year at the site where trees were felled soon after the flowering period. We argue that when male floral resources are altered in exploited populations, pollinators of generalist nature may show a shift in foraging pattern. Also, generalist plants may be as susceptible to pollination loss as are specialist plants. The need for outcrossing in the species would probably exacerbate this vulnerability.     

Gender specialization and factors affecting fruit set of the wind-pollinated heterodichogamous Juglans regia

Gender expression, flowering phenology, reproductive performance and factors affecting fruit set (i.e., flowering synchrony, size and distance to the nearest pollen donor) were investigated in a cultivated population of a wind-pollinated self-compatible heterodichogamous Juglans regia (Juglandaceae). Four flowering morphs, (i.e., protandrous, protogynous, male and female) were observed. The sexual functions of the protandrous and protogynous morphs were almost synchronous; however, they were not reciprocal, and the separation of male and female flowering within most monoecious individuals was not complete. Thus, within-morph mating and geitonogamous pollination may be common. The ratio of protandrous versus protogynous morphs was biased towards the protandrous morph, but the fruit set did not differ between the morphs, suggesting that the fruit set of the protandrous morph could be partly compensated by within-morph pollination. The ratio of the female flower number or fruit number to the total male catkin length was higher in the protogynous morph than in the protandrous morph and did not vary with plant size, suggesting that gender variation was not size dependent and that the sexual function of protandrous morphs was more male biased. Fruit set depended on plant size only for protandrous morphs. The fruit set of individual plants decreased with increasing distance to the nearest pollen donor regardless of morph, possibly because of pollen limitation. The fruit set of individual plants increased with flowering synchrony, indicating that flowering synchrony could affect reproductive success.     

A multi-year study of factors affecting fruit production in Aristolochia paucinervis (Aristolochiaceae)

Pollen limitation, resource limitation, fruit abortion, and predation have all been proposed as factors explaining low fruit set in hermaphroditic plants. We conducted a 5-year study combining field observations and pollination experiments to determine the causes of the low fruit set in Aristolochia paucinervis, a Mediterranean species with a specialized pollination system in two populations in SW Spain. Fruit initiation was markedly low, and between 28.6 and 75.0% of the flowering stems did not initiate any fruit. In most flowers, the number of germinated pollen grains was less than the number of ovules, and supplemental pollination significantly increased fruiting, indicating deficient pollination. In A. paucinervis, autonomous self-pollination seems to be a decisive factor in fruit production because the number of germinated pollen and the fruit set from flowers bagged before anthesis were similar to those in free-pollinated flowers. Only in 2005 did flowers that were successfully pollinated outnumber ripened fruits, suggesting that other factors limit fruiting. We found a significant positive correlation between tuber mass and fruit set. Deficient pollination and lack of resources could explain the low fruit set, but the relative consequences seem to vary spatially and temporarily.     

Apical pattern of fruit production in the racemes of Ceratonia siliqua (Leguminosae: Caesalpinioideae): role of pollinators

Fruit production and arrangement within the raceme were studied in two dioecious populations of Ceratonia siliqua (Leguminosae: Caesalpinioideae), an arboreal species that produces caulogenous racemes (emerging only from the old branches) with numerous flowers. Fruit production per raceme was low and similar between years and populations and even between individuals. During flowering, there were considerable flower losses from predation and lack of pollination. A mean of nine flowers per raceme began the transformation into fruits, of which 77% aborted. The final fruit production per raceme increased significantly following hand pollination, but was always very much lower than the availability of flowers in the raceme. The results suggest that fruit production of each raceme is limited by both availability of resources and a deficient pollination. In racemes setting fruit arrangement follows a definite pattern that remains constant between years and populations: fruit production was significantly higher in the apical zone of the raceme and lower in the basal zone. The pollinators of C. siliqua (flies and wasps) showed a clear preference for beginning their visits at the apex of a raceme. As a result, the pollen load deposited on the stigmas decreased from apex to base of the raceme. In most of the flowers situated in the central and basal zone of the raceme, the number of pollen grains deposited on their stigmas was lower than the number of their ovules. The high number of seeds in developed fruits suggests that the plant selectively aborts flowers that receive a smaller pollen load. The results indicate that the final pattern of fruit arrangement within the raceme is a direct result of pollinator activity.     

Abscisic Acid and cutout in cotton

A decline in growth, flowering, and boll (fruit) retention is referred to as cutout in cotton (Gossypium hirsutum L.). Fruit load affects cutout, possibly through hormonal effects. Experiments were conducted to test the hypothesis that fruits are a source of abscisic acid (ABA) that moves into fruiting branches and growing points where it inhibits growth, flowering, and boll retention. Removal of the flower or young boll at the first node of fruiting branches did not decrease the ABA content of fruiting branches or the abscission zone at the second node. Effects on ABA content of the boll at the second node varied. In one field test, ABA content of bolls at the second node decreased with successive harvests as bolls were removed from first node positions of several fruiting branches. Thus, the effect was cumulative and was not limited to individual branches. Removal of the flower or boll at the first node increased boll retention at the second node. Removal of all flowers during the first 3 weeks of flowering delayed the decreases in growth, flowering, and boll retention that occurred as fruit load increased. But, the ABA content of fruiting branches and mainstem apices was not decreased by early defruiting and did not increase with increasing fruit load. The results do not support the hypothesis that fruits are a source of ABA that moves into fruiting branches and growing points where it then inhibits growth, flowering, and boll retention.     

Influence of pollination systems on fruit set and fruit quality in kiwifruit (Actinidia deliciosa)

Kiwifruit (Actinidia deliciosa (Chev.) Liang and Ferguson) is a dioecious species highly pollination dependent, since fruit size is directly correlated with seed number. To evaluate the best pollination system for this species a study of two natural and two artificial pollination systems was carried out and their effect on fruit set and fruit quality was determined. While anemophilous pollination produced few and small fruits, that were not commercially acceptable, high fruit set percentages and good quality fruits were obtained when insects participated in the pollination process. Artificial pollination, performed as hand pollination, improved yield quantity and quality. When environmental conditions impair insect activity this method can be an useful agricultural practice in order to assure a good yield.     

优质荔枝新品种'草莓荔'开花与落果特性观察

从植株群体和单穗调查分析‘草莓荔’雌雄花开花特性,结果表明,‘草莓荔’雌雄花交替开放的顺序主要是雌→雄,即前期开雌花,后期开雄花;但也有少量交替开放顺序是雄→雌→雄,即先开雄花,中期开雌花后再开雄花。植株群体的花期和雌雄花开放相遇时间比单穗长,更有利于传粉受精。落果从花后15d开始出现,并在花后18～25d出现第一次落果高峰,落果最多,开花后51～58d为第二次落果高峰。     

ECOLOGICAL CORRELATES OF REPRODUCTIVE TRAITS OF MEXICAN RAIN FOREST TREES

Sexual systems of 139 tree species from a tropical rain forest at Los Tuxtlas, Mexico were investigated to: 1) estimate the relative proportions of hermaphroditic, monoecious, and dioecious species; 2) describe flowers, fruits, and seeds in terms of size and weight; 3) describe flowering and fruiting phenology; and 4) correlate sexuality to pollination and dispersal syndromes, and the successional status occupied in the forest. Hermaphroditism occurred in 63% of the species, monoecism in 9%, and dioecy in 27%. Nondioecious species had larger flowers, but dioecious species had more seeds per fruit. The dioecious condition was associated with small flowers pollinated with unspecialized insects and fleshy fruits dispersed by animals at both species and generic levels. Reproductive traits were more correlated among nondioecious species than dioecious species. Pioneer species had more seeds per fruit, and longer flowering and fruiting periods, but persistent species produced heavier seeds and fruits. Flower and fruit morphological traits, sexual systems, and tree guilds are related in a comprehensive way, and a flow model based on data from this study is presented.     

Do consecutive flower visits within a crown diminish fruit set in mass‐flowering Hancornia speciosa (Apocynaceae)?

Hancornia speciosa is a self-incompatible, mass-flowering, sphingophilous fruit crop (mangaba) of northeast and central Brazil. The flowers have a precise pollination apparatus, which optimizes pollen transfer between flower and pollinator. While the pollination mechanism avoids self-pollination, mass-flowering promotes geitonogamy. During a flower visit, almost half of the exogenous pollen grains adhering to the proboscis are deposited on the stigma surface. A pollination experiment with a nylon thread simulating six consecutive flower visits within a crown revealed that only the first two flowers visited (positions 1 and 2) are highly likely to set fruit. Super-production of flowers, and consequently obligate low fruit set, seem to be part of the reproductive strategy of the obligate outcrossing plant, Hancornia speciosa.     

Reproductive Traits and Phenology of Plants in Tropical Dry Evergreen Forest on the Coromandel Coast of India

Qualitative reproductive traits of 84 plant species belonging to 41 families were studied in tropical dry evergreen forest on the Coromandel coast of India. Majority of species had rotate type, white-coloured, scented flowers, rewarding nectar and pollen and pollinated chiefly by bees. An association between floral traits and pollination spectrum is evident. Bee pollination was prevalent in pollination systems. Among the fruit types, drupe and berry were common in black and red colour respectively, and dispersed by zoochorous mode. Seeds of brown- and green-coloured dry fruits, without any reward were disseminated by wind and explosion. The reproductive phenophase of trees and lianas occurred mostly during the dry period from January to June, which receives rainfall of less than 50 mm a month. However, shrubs showed a peak in flowering and fruiting in wet period. Detailed phenological observations of 22 woody species revealed a seasonal and unimodal pattern in flowering. Although some species were in flower round the year, flowering activity was skewed towards the dry season. The fruiting activity showed a bimodal pattern, one peak in dry season and another in wet season. Many species displayed a temporal aggregation in flowering and fruiting. The significant relation was obtained between reproductive traits and phenology of plants in the tropical dry evergreen forest.     

优质荔枝新品种‘草莓荔’开花与落果特性观察

从植株群体和单穗调查分析‘草莓荔’雌雄花开花特性,结果表明,‘草莓荔’雌雄花交替开放的顺序主要是雌→雄,即前期开雌花,后期开雄花;但也有少量交替开放顺序是雄→雌→雄,即先开雄花,中期开雌花后再开雄花。植株群体的花期和雌雄花开放相遇时间比单穗长,更有利于传粉受精。落果从花后15d开始出现,并在花后18～25d出现第一次落果高峰,落果最多,开花后51～58d为第二次落果高峰。     

文冠果人工种群的果实发育与落花落果特性研究

以甘肃定西生长的文冠果为对象,探讨了文冠果果实发育特征及落花落果规律。结果表明：（1）文冠果果实生长发育历经约2.5个月,生长发育主要在花期后的40 d内。果实发育特点为：在花期过后的10 d内生长最快,增长率达193%,此后一月仍保持较快的生长速度,月增长率达120%,而最后一月生长比较缓慢,月增长率仅为7%,整个发育期内横径发育速度一直大于纵径;（2）文冠果落花落果主要发生在果实生长发育的初期,且落花落果率高,达94.65%(落花率5.74%,落果率88.91%）,坐果率低,只有5.35%。其落果特点为：落果是一个持续的过程,自花期过后果实膨大到成熟一直在落果,但也存在一个落果的高峰期,即花期过后的10 d内,落果最严重,落果率约达40%。研究认为,从保花保果的角度考虑,开花末期是文冠果水肥管理的重点时期。但从文冠果开花过于繁茂和可孕花分布过于集中而大量消耗树体和顶花序养分,从而造成严重的落花落果的现实考虑,应在开花末期和幼果期疏花疏果或任其自然落花落果,但均应加强果实发育中后期的水肥管理,从而提高坐果率。     

Perfect syncarpy in apple (Malus x domestica 'Summerland McIntosh') and its implications for pollination, seed distribution and fruit production (Rosaceae: Maloideae)

BACKGROUND AND AIMS: The gynoecium of the domestic apple, Malus x domestica, has been assumed to be imperfectly syncarpic, whereby pollination of each stigmatic surface can result in fertilization within only one of the five carpels. Despite its implied effect on fruit quantity and quality, the resulting influence of flower form on seed set and distribution within the apple fruit has seldom been investigated. Instead, poor fruit quality is usually attributed to problems with pollination, such as low bee numbers and/or ineffective pollinators within apple agro-ecosystems. The objective of this study was to determine the true nature of gynoecial structure and its influence on fruit production in the apple cultivar 'Summerland McIntosh'. METHODS: A stigma-excision method was used to determine the effects of uneven pollination among the five stigmas on fruit quantity (as measured by fruit set), and quality (seed number and distribution). In addition, flowers were examined microscopically to determine pollen tube pathways. KEY RESULTS: Fruit set, seed number, seed distribution, and the microscopic examination of flower gynoecial structure reported in this study indicated that the gynoecium of the cultivar Summerland McIntosh is perfectly syncarpic and not imperfectly syncarpic as previously thought. CONCLUSIONS: Pollination levels among the five stigmas need not be uniform to obtain full seed development within Summerland McIntosh fruit; even if one stigmatic surface is adequately pollinated, a full complement of seeds is likely. The importance of perfect syncarpy in recognizing true causes of poor fruit quality in apple is discussed.