Causes of spatial patterns of fruit set in waratah: Temporal vs. spatial interactions between flowers on an inflorescence

Spatial patterns of fruit set within inflorescences may be controlled by pollination, nutrient allocation, or inflorescence architecture. Generally, flowers that have spatial and/or temporal precedence are more likely to set fruits. We sought to separate these factors by comparing patterns of fruit set on inflorescences of two species of Telopea (Proteaceae); one that flowers from the tip to the base of the rachis, the other from base to tip. In both species, most fruits were set at the top of the inflorescence (the last flowers to open for T. speciosissima) and this was extreme for T. mongaensis, where the top flowers open first. Fruit set was not generally limited by inadequate pollination for either T. mongaensis or T. speciosissima, as hand pollinations did not increase fruit set and many abscised flowers contained pollen tubes. In T. speciosissima, we tested whether removal of developing topmost fruits would ‘release’ those that had initiated but not yet aborted lower down. There was no significant effect. Plant hormones can increase the degree to which a developing fruit is a sink for nutrients, so we applied cytokinin to the developing lower fruits on some inflorescences. There was no significant effect of the hormone treatment. We conclude that temporal precedence may contribute to the skewed pattern of fruit set in T. mongaensis, because there was an extreme concentration of fruit set on the distal part of the inflorescences, but it cannot explain this pattern of fruit set in T. speciosissima, where the distal flowers are the last to open. Some other process must therefore constrain fruit set to the topmost flowers in an inflorescence. While cytokinin application had no significant effect, the power of this experiment was low and we consider that the hypothesis of hormonal control is worth further exploration.     

荒漠植物准噶尔无叶豆结实、结籽格局及其生态适应意义

连续2a采用野外记录的方式对荒漠植物准噶尔无叶豆的结实、结籽格局进行了比较研究。结果表明:植株内花序生成格局表现为不同大小的花序在总花序中的比重与其结实率存在显著正相关,其中含有中等花数目的花序所占的比重及其结实率均最高,此种分布格局能够最大程度的保证繁殖成功;果序内果实生成格局表现为单花着生位置(从近柄端算起)与其结实量占总结实量的比例成线性负相关,花序基部的结实比例最高,顶部结实比例近为0;荚果内种子生成格局表现为:中间位置的胚珠败育率最低。另外,年际间荚果内仅有(1.08±0.03)粒和(1.07±0.03)粒种子能够完全成熟,此种结籽格局是准噶尔无叶豆保证后代质量的最佳策略。     

Analysis of the tomato fruit growth response to temperature and plant fruit load in relation to cell division, cell expansion and DNA endoreduplication

BACKGROUND AND AIMS: To better understand the regulation of fruit growth in response to environmental factors, the effects of temperature and plant fruit load on cell number, cell size and DNA endoreduplication were analysed. METHODS: Plants were grown at 20/20 degrees C, 25/25 degrees C and 25/20 degrees C day/night temperatures, and inflorescences were pruned to two ('2F') or five ('5F') flowers. KEY RESULTS AND CONCLUSIONS: Despite a lower fruit growth rate at 20/20 degrees C, temperature did not affect final fruit size because of the compensation between cell number and size. The higher cell number at 20/20 degrees C (9.0 x 10(6) against 7.9 x 10(6) at 25/25 degrees C and 7.7 x 10(6) at 25/20 degrees C) resulted from an extended period of cell division, and the smaller cell size was due to a shorter period of expansion rather than a lower expansion rate. By contrast, the lower fruit growth rate and size of 5F fruits compared with 2F fruits resulted from the slow down of cell expansion, whereas the number of cells was hardly affected in the proximal fruit. However, within the inflorescence the decreasing gradient of fruit size from proximal to distal fruits was due to a decrease in cell number with similar cell size. Fruit size variations within each treatment were always positively correlated to variations in cell number, but not in cell size. Negative correlations between cell size and cell number suggested that cells of tomato pericarp can be seen as a population of competing sinks. Mean ploidy was slightly delayed and reduced in 5F fruits compared with 2F fruits. It was highest at 25/25 degrees C and lowest at 25/20 degrees C. Treatments did not affect ploidy and cell size in similar ways, but within each treatment, positive correlations existed between mean ploidy and cell size, though significant only in the 2F-25/20 treatment.     

Factors constraining fruit set in Mandevilla pentlandiana (Apocynaceae)

The reproductive success of Mandevilla pentlandiana was studied to disclose its reproductive strategy, and to determine the links between nectar production, breeding system, fruit set and inflorescence size. The plant produces many inflorescences with a large number of flowers but initiates few fruits (9%). This vine is self-compatible but not autogamous. Given that no significant differences could be detected considering many traits (ripe and abortive fruit sets, fruit quality, and seedling survival) between the pollination treatments (self-, cross-and natural-), the low natural fruit set was not related to pollen limitation. Fruits were not distributed at random within inflorescences (earlier fruits had the highest probability of maturation) but there were no significant differences in fruit quality according to different fruit positions. Conversely, the time of fruit initiation influenced most of the fruit-traits. Many developing fruits were aborted (20%). An increase in the probability of abortion was detected when the whole inflorescence was hand pollinated. In addition, a positive correlation was detected between the abortions and the number of ripe fruits which developed before them. Looking at our data from an evolutionary perspective, we argue that a theoretical inflorescence size, corresponding to the intersection point between the mean values of fruit number and fruit set per inflorescence, can be assumed to indicate the optimum inflorescence size that maximizes equally both female and male functions. Comparison between the theoretical and the observed mean inflorescence size suggests, that for M. pentlandiana , pollen donation may be the primary evolutionary factor behind excess flowers.     

Sequential decline in fruit resource allocation within inflorescences of Sagittaria trifolia: a test of non‐uniform pollination hypothesis

Flowering plants often exhibit declining resource investment to floral organs, fruits and seeds temporally or spatially in an inflorescence. To account for such variances, non‐uniform pollination hypothesis, which highlights various mating environments each flower experiences, provides adaptive significance for allocation patterns but with controversial supports. Sagittaria trifolia (Alismataceae) was used to examine differences in seed number, seed weight and germination rate among sequential fruits within inflorescences. Ovule number was also investigated to evaluate allocation patterns in the floral stage. To test the non‐uniform pollination hypothesis, we used three polymorphic microsatellite loci of S. trifolia to estimate the seed outcrossing rate in proximal and distal fruits. The results showed that the seed number, average seed weight and seed germination rate of S. trifolia gradually decreased from proximal to distal fruits within inflorescences. The percent of decrease in seed number between two contiguous fruits was 14.68 ± 3.22%, which was much stronger than the percent of decrease in ovule number at 6.95 ± 1.60%. Both proximal and distal fruits within inflorescences had high outcrossing rates (81.5 ± 5.0%, proximal; 82.3 ± 6.9%, distal) and they did not differ significantly. Overall, there was an acropetal decline of resource allocation to fruits within inflorescences of S. trifolia. Allocation pattern to ovules was not a limiting factor for seed production. The lack of difference in outcrossing rate between proximal and distal fruits indicated that the allocation strategy was probably not caused by non‐uniform pollination, but more likely position effects.     

Fruit size decline from the margin to the center of capitula is the result of resource competition and architectural constraints

Plants produce repeated structures, such as leaves, flowers, and fruits, which differ in size and shape. One example of this is fruit size, which is commonly observed to decrease from proximal to distal positions within an inflorescence. The resource limitation hypothesis proposes that because proximal fruits usually develop first, they have temporal priority on access to resources over distal fruits. The non-mutually exclusive architectural effects hypothesis suggests that these position effects in fruit size may also be due to inherent architectural variation along infructescence axes. We separated out the effects of resource competition and inflorescence architecture by removing the outer or the inner flowers within capitula of Tragopogon porrifolius. We also studied if fruit position influenced germination and seedling performance in order to assess fitness consequences of position effects. Inner fruits were significantly heavier when outer flowers were removed. However, outer fruits did not significantly increase when inner flowers were removed, suggesting later fruits were limited by the development of early fruits. Our findings also suggest that architectural constraints restricted the size of inner fruits in comparison with outer ones. We found that both resource competition and inflorescence architecture affected the fruit size of T. porrifolius, even though this species does not have linear, indeterminate inflorescences. We advance the hypothesis that, when such effects on fitness occur, resource competition-mediated position effects could turn, in evolutionary time, into architectural position effects.     

Quantifying sink and source limitations on dry matter partitioning to fruit growth in peach trees

We describe an approach for determining the degree of sink and source limitations on peach ( Prunus persica L. Batsch) fruit growth during several growth periods. Source limitations on fruit growth may be due to either a shortfall in assimilate supply within the tree (supply limitation) or to a deficiency in the capacity of the translocation system to deliver assimilates in sufficient quantity to support the maximum fruit growth rate (transport/competition limitation). To ascertain the potential maximum rate of fruit growth, fruit thinning treatments were used. One month after bloom, the number of fruits per tree was adjusted to between 50 and 700 on an early and a late maturing peach cultivar (cvs Spring Lady and Cal Red, respectively). Rates of potential sink demand, potential source supply and actual fruit growth were estimated from sequential harvests of all fruits on 42 trees on two (Spring Lady) and three (Cal Red) dates. These values were used to estimate the proportion of potential growth achieved, and the supply and transport/competition limitations on fruit growth. The results indicated that source limitations were significant on trees with moderate to high fruit numbers. These source limitations were due to supply limitations during all harvest intervals and to transport/competition limitations during the early harvest intervals. Sink limitations occurred to the greatest extent during the mid-period of fruit growth on the later maturing cultivar.     

Variation in fruit- and seed set among and within inflorescences of Melampyrum roseum var. japonicum (Scrophulariaceae)

Abstract Variations in fruit set and seed set among and within inflorescences of the annual herb, Melampyrum roseum var. japonicum, were studied. Under natural conditions, although the mean fruit set was slightly different among inflorescences, the mean seedset was not significantly different among inflorescences within the plants. In constrast, within the inflorescences, the flowers located at a lower position of the inflorescence and which opened earlier showed higher fruit set than those at a higher position and which opened later. However, the seed set of matured capsules were not significantly different from each other, regardless of the position of flowers within the inflorescences. Patterns of the fruit- and seed set under open pollination indicated that variation in seed reproduction of M. roseum is due to variation in fruit production. The results of clipping experiments of flowers revealed that there was no functional limitation in seed production among flowers located at various positions within the inflorescence. It seemed that the variation in the fruit set within the inflorescences of M. roseum was not attributable to 'architectural effects'. Reduction of the number of flowers within the inflorescences resulted in an increase of fruit set and seed weight, indicating that the flowers in an inflorescence compete for resources. This phenomenon supports the 'resource competition hypothesis', and variation in fruit set within the inflorescence is attributable to competition among flowers within the inflorescence for limited resources. Consequently, it was concluded that, under natural conditions, the early blooming flowers located at lower positions of the inflorescences obtain more resources and produce more fruits than the late blooming flowers located at higher positions in M. roseum .     

Effects of fruit set sequence and defoliation on cell number,cell size and hormone levels of tomato fruits (Lycopersicon esculentum Mill.) within a truss

Fruit size within a tomato (Lycopersicon esculentum Mill.) truss depends on both fruit position in the truss and the time of pollination among fruits. In the natural pollination sequence a difference of 5 days in the pollination of proximal and distal flowers results in significant final size differences between proximal and distal fruits. These final size differences were eliminated when all flowers were pollinated simultaneously. At anthesis proximal ovaries have higher cell numbers than distal ovaries but the cell division activity and cell enlargement in both positions was similar in the first 10 days of fruit growth. Simultaneous pollination resulted in lower cell numbers in proximal but higher cell numbers in distal fruits compared to control fruits.Hormone levels in different sized fruits were measured using radioimmunoassays. Cytokinin concentration during the cell division period indicated a possible role in the regulation of cell division. With other hormones no obvious correlations were found. The results are discussed in relation to factors determining final fruit size in tomato.     

The effect of assimilate supply on fruit growth and hormone levels in tomato plants

Rates of dry matter accumulation and contents of starch, sugars and abscisic acid (ABA) of tomato fruits differed significantly during development at three positions (proximal, middle and distal) on a truss. Proximal fruits, which accumulated dry matter most rapidly during early development, generally had least ABA (per g DW).Partial defoliation reduced carbon accumulation by all fruits but increased ABA, especially in distal fruit, and indoleacetic acid (IAA), particularly in proximal fruits. The ABA content of leaves in partially defoliated plants was similar to that of leaves on non-defoliated plants.Removal of distal fruits on a truss enhanced carbon movement to the remaining proximal fruits and also increased their ABA content early in development but did not affect their IAA content. On the other hand, when proximal fruit were removed there was no large or lasting increased accumulation of carbon by the remaining distal fruits and they contained less ABA and IAA than fruits on plants without fruit thinning. Leaf carbon and ABA levels showed no marked trend in response to fruit thinning.The amount of carbon in the stems was increased by fruit thinning but decreased by partial defoliation.The possible roles of ABA and IAA in regulating fruit growth are discussed.Part of this work has been presented to a Symposium on Phloem loading and related processes at Bad Grund/Oberhar, W. Germany, July 1979.     

A rapid increase in cytokinin levels and enhanced ethylene evolution precede Al3+-induced inhibition of root growth in bean seedlings (Phaseolus vulgaris L.)

Fruit sink strength or its ability to attract assimilates depends bothon sink activity and size. This study investigated one main component of sinksize, that is the number of fruit cells during tomato plant development. Plantswere grown in a controlled climate chamber under a limiting (LS, six fruits pertruss) and non-limiting (NLS, two fruits per truss and CO₂enrichment) supply of carbon assimilates. Under NLS conditions, fruit cellnumber was homogeneous among successive trusses, and fruits contained onaverage1.2×10⁶ more cells than under LS conditions,though differences were not significant on the first truss which underwent thelowest competition. Under LS conditions, an ontogenetic increase in cell numberwas observed in proximal fruits of the upper trusses attributed to theenlargement of the apical meristem during plant development. The decrease ofcell number from proximal to distal fruits within a truss, that was expectedfrom the literature, was generally observed in the LS experiment, with anaverage significant difference of about1.6×10⁶cells between the first and fifth fruits. Nevertheless, whereas the gradient incell number from proximal to distal fruits was steep in the upper trusses, itwas not significant on the lower trusses indicating that this gradient largelydepended on the level of competition during floral development. Thus, under lowassimilate supply, cell division is a main limiting factor for fruit growth,although cell enlargement during further fruit development is also affected,butwas not measured in this work.     

Effects of natural rates of geitonogamy on fruit set in Asclepias speciosa (Apocynaceae): evidence favoring the plant's dilemma

The role of geitonogamy in the evolution of inflorescence design is not well understood. The plant's dilemma hypothesis proposes that evolution of larger inflorescences is driven by selection for greater pollinator attraction, but constrained by higher rates of geitonogamy experienced by larger inflorescences. Here we investigate the role of geitonogamy on fruit set in natural populations of Asclepias speciosa. We compared fruit set from three pollination treatments: (1) inflorescences bagged before and after receiving 6 hand outcross pollinia (Bag), (2) inflorescences unbagged and receiving 6 hand outcross pollinia (Open), and (3) naturally pollinated inflorescences (Control). The Bag and Open treatments initiated significantly more fruits than the Control. Bag aborted significantly fewer fruits than Open or Control. Fruit set was significantly higher in Bag than Open, and Open had significantly higher fruit set than Control. From these results, we conclude that (1) high rates of geitonogamy significantly increase fruit abortion and reduce fruit set in natural populations of A. speciosa and (2) natural populations are compatible pollen limited. Both findings are consistent with the plant's dilemma hypothesis.     

Modulation of Competition between Fruits and Leaves by Flower Pruning and Water Fogging, and Consequences on Tomato Leaf and Fruit Growth

The effects of water fogging and reducing plant fruit load werestudied in a tomato crop grown in a glasshouse under Mediterraneansummer conditions. The objective of these treatments was toreduce competition between leaves and fruits for carbohydratesand water. Flower pruning increased plant leaf area and increasedfruit, stem, lamina and petiole dry mass (DM). This indicatesthat leaf area growth was limited during the summer due to competitionbetween fruits and leaves for assimilates. In contrast, reducingthe air vapour pressure deficit (VPD) by water fogging had noeffect on plant leaf area or aerial plant DM. Interestingly,there was a significant interaction between plant fruit loadand VPD: the higher the leaf[ratio]fruit ratio the greater theresponses to a reduction in VPD (increase in fruit DM, fruitdiameter, fruit and leaf expansion rate). The data suggest thatunder high fruit loads, water and carbohydrates limit growthunder Mediterranean summer conditions. However, reducing VPDwas not always sufficient to enhance fruit and leaf growth.This might be due to the lower leaf area under high fruit load.In contrast, reducing VPD under low fruit load triggered higherrates of leaf and fruit expansion; this is probably linked toa greater availability of water and carbohydrates. Copyright2001 Annals of Botany Company Assimilate competition, assimilate supply, flower pruning, fruit load, fruit growth, generative/vegetative growth, leaf growth, Lycopersicon esculentum, specific leaf weight, tomato, vapour pressure deficit, water stress     

Evolution of growth habit, inflorescence architecture, flower size, and fruit type in Rubiaceae: its ecological and evolutionary implications

During angiosperm evolution, innovations in vegetative and reproductive organs have resulted in tremendous morphological diversity, which has played a crucial role in the ecological success of flowering plants. Morindeae (Rubiaceae) display considerable diversity in growth form, inflorescence architecture, flower size, and fruit type. Lianescent habit, head inflorescence, small flower, and multiple fruit are the predominant states, but arborescent habit, non-headed inflorescence, large flower, and simple fruit states occur in various genera. This makes Morindeae an ideal model for exploring the evolutionary appearances and transitions between the states of these characters. We reconstructed ancestral states for these four traits using a bayesian approach and combined nuclear/chloroplast data for 61 Morindeae species. The aim was to test three hypotheses: 1) self-supporting habit is generally ancestral in clades comprising both lianescent and arborescent species; 2) changes from lianescent to arborescent habit are uncommon due to "a high degree of specialization and developmental burden"; 3) head inflorescences and multiple fruits in Morindeae evolved from non-headed inflorescences and simple fruits, respectively. Lianescent habit, head inflorescence, large flower, and multiple fruit are inferred for Morindeae, making arborescent habit, non-headed inflorescence, small flower, and simple fruit derived within the tribe. The rate of change from lianescent to arborescent habit is much higher than the reverse change. Therefore, evolutionary changes between lianescent and arborescent forms can be reversible, and their frequency and trends vary between groups. Moreover, these changes are partly attributed to a scarcity of host trees for climbing plants in more open habitats. Changes from large to small flowers might have been driven by shifts to pollinators with progressively shorter proboscis, which are associated with shifts in breeding systems towards dioecy. A single origin of dioecy from hermaphroditism is supported. Finally, we report evolutionary changes from headed to non-headed inflorescences and multiple to simple fruits.     

Within-bunch Variability in Banana Fruit Weight: Importance of Developmental Lag Between Fruits

Within-bunch (inflorescence) variability in banana fruit weightis of great importance: distal fruits (at the bottom of thebunch) are 30 to 40% smaller than basal fruits at the top. Wehypothesize that this variability is related to a developmentallag between fruits. To validate this hypothesis, histologicalstudies (evolution in number of cells along the fruit radius,starch granule number and size) associated with physiologicalmeasurements (pulp dry weight, dry matter and starch concentration)were carried out. Fruit development stages were dated in cumulativedegree-days (dd) from flower emergence to 3 weeks after theharvest stage (1300 dd). For a fruit located at the top of thebunch, cell divisions ceased around 350 dd and cells began tofill with starch as soon as they appeared. A developmental lagbetween fruits at the top and bottom of the bunch was observed:cell divisions started and stopped approx. 70 dd later in bottom(distal) compared to top (basal) fruits. At the end of celldivisions, basal fruits had a higher number of cells along thefruit radius. This difference in cell number may be due to increasedcompetition for assimilates between fruits when cell divisionoccurs in distal fruits. Variability in cell number may be relatedto variability in pulp dry weight. We conclude that within-bunchvariability in banana fruit weight is related to a differencein cell number and age. Copyright 2001 Annals of Botany Company Musa acuminata, banana, fruit development, fruit growth, cell number, starch accumulation, fruit quality, fruit green-life, fruit-fruit competition     

Selective fruit abortion in Prunus mahaleb (Rosaceae)

In Prunus mahaleb (Rosaceae), only a small proportion of those flowers that initially develop give rise to fruits, and the probability of producing fruit differs between flowers. The results of this study indicate that the probability of producing fruit is about 60% for the first (i.e., most proximal) flower to develop in each inflorescence, but very low for the last (i.e., most distal). This pattern is maintained regardless of between-inflorescence differences in flowering phenology, suggesting that each inflorescence can be viewed as an independent subunit with respect to allocation of resources for reproduction. These results are compatible with the ovary reserve and selective abortion hypotheses for the adaptive value of low fruit set in P. mahaleb.     

A multilevel analysis of fruit growth of two tomato cultivars in response to fruit temperature

Fruit phenotype is a resultant of inherent genetic potential in interaction with impact of environment experienced during crop and fruit growth. The aim of this study was to analyze the genetic and physiological basis for the difference in fruit size between a small (‘Brioso’) and intermediate (‘Cappricia’) sized tomato cultivar exposed to different fruit temperatures. It was hypothesized that fruit heating enhances expression of cell cycle and expansion genes, rates of carbon import, cell division and expansion, and shortens growth duration, whereas increase in cell number intensifies competition for assimilates among cells. Unlike previous studies in which whole‐plant and fruit responses cannot be separated, we investigated the temperature response by varying fruit temperature using climate‐controlled cuvettes, while keeping plant temperature the same. Fruit phenotype was assessed at different levels of aggregation (whole fruit, cell and gene) between anthesis and breaker stage. We showed that: (1) final fruit fresh weight was larger in ‘Cappricia’ owing to more and larger pericarp cells, (2) heated fruits were smaller because their mesocarp cells were smaller than those of control fruits and (3) no significant differences in pericarp carbohydrate concentration were detected between heated and control fruits nor between cultivars at breaker stage. At the gene level, expression of cell division promoters (CDKB2, CycA1 and E2Fe‐like) was higher while that of the inhibitory fw2.2 was lower in ‘Cappricia’. Fruit heating increased expression of fw2.2 and three cell division promoters (CDKB1, CDKB2 and CycA1). Expression of cell expansion genes did not corroborate cell size observations.     

Variability in space and time: contrasting fruit distribution patterns in the deceptive orchid Orchis militaris

• In angiosperms, a decrease in fruit production towards the apex of individual inflorescences is usually observed. Orchids are thought to be primarily pollination‐limited species, and non‐uniform pollination could cause this decrease pattern in several species. Fruit production was investigated in relation to flower position and floral display size in Orchis militaris (Orchidaceae), a deceptive species.
• Over 2 years, eight populations of O. militaris were studied and fruit position along the inflorescence was recorded. Generalised linear models were performed to examine the effect of population, year, flower position and floral display size on fruit production.
• The dominant pattern was characterised by a higher fruit set in the middle part of the inflorescence (parabolic pattern). A non‐directional pattern of fruit production was also detected in some populations. Within a given population, patterns were generally consistent among years. In one of the two study years and in one of the eight populations specifically, the proximal‐to‐distal decrease in fruit production was dramatic in plants with a large floral display but weak or absent in small displays.
• Our study demonstrates the intraspecific diversity of fruit distribution patterns in O. militaris. Non‐uniform pollination along the inflorescence is likely to be responsible for the parabolic pattern, while irregular visitation could explain the non‐directional pattern of fruit production. Pattern variation among years and between populations could arise from spatiotemporal variation in pollinator assemblages. Resource competition effects could explain the interaction effect between display size and flower position.

     

The Regulatory Role of Inflorescence Leaves in Fruit-setting by Sweet Orange (Citrus sinensis)

Inflorescence leaves improve fruit set on sweet orange trees. We sought an explanation for this effect in terms of carbon demand by developing fruit versus potential supply from adjacent leaves. Our assessment was based upon measurements of fruit growth, leaf photosynthesis and ¹⁴C distribution patterns in plants grown under controlled conditions. Leafy inflorescences had sufficient foliar surface (1.24 dm²) and photosynthetic capacity (CO₂ 10.1 mg · dm^-2· h^-1) to support early development of fruits on the same shoot, and to make a substantial contribution towards subsequent growth. ¹⁴C-assimilates derived from new leaves were distributed towards adjacent fruit which showed strong competition for labelled substrate. By contrast, fruit borne on leafless inflorescences had to obtain all their assimilates from older leaves whose photosynthetic capacity (CO₂ 3.5–4.6 mg · dm^-2· h^-1) and individual area (0.2 dm²) were generally insufficient to wholly sustain fruit growth, so that a large number of old-leaves were needed; these fruit would be more susceptible to competition from other sinks.