Sink strength manipulation in branches of a Mediterranean woody plant suggests sink-driven allocation of biomass in fruits but not of nutrients in seeds

The relative autonomy of branches within an individual plant may favor different resource allocation responses after reproductive losses. The assessment of these reproductive strategies at the branch level before their integration at the plant level would provide more insight into how plants deal with reproductive losses. Here, we present a field experiment to assess changes in the allocation strategies at the branch level after sink strength manipulation in a Mediterranean woody plant (Cistus ladanifer). We applied three levels of removal of developing fruits (75, 25 and 0 %) on branches of the same plants, and measured their effects on resource allocation (biomass, nitrogen and phosphorus) and seed production after controlling for the effects of branch diameter and leaf weight per branch. Our results suggest that after experimental fruit thinning, C. ladanifer branches presented a sink-driven allocation of biomass to fruits but this was not the case for the allocation of nutrients to seeds, which could be driven by competition with leaf biomass. Reductions in biomass per fruit resulted in a reduction in seed output since the average weight per seed remained constant. From these results, it could be suggested that an heterogeneous distribution of fruit losses among the branches within a crown would produce a higher impact on reproductive output than a more equitable distribution.     

Shoot Apex Demand Determines Assimilate and Nutrients Partitioning and Nutrient-uptake Rate in Tobacco Plants

Our previous experiment revealed that apex-removed plants have larger root systems but a lower K＋-uptake rates than intact tobacco plants. Since the apex is not only a center of growth and metabolism, but also an important place of auxin synthesis and export, the aims of this study were to distinguish whether the apex demand or auxin synthesized in the apex regulates assimilate and nutrients partitioning within plant, and to explain the reason for the lower K＋-uptake rate of the apex-removed plant. In comparison with the control plant, covering the shoot apex with a black transparent plastic bag reduced net increases in dry matter and nutrients; however, the distribution of the dry matter and nutrients between shoot and roots and nutrient-uptake rates were not changed. Removal of the shoot apex shifted the dry mass and nutrients distributions to roots, and reduced the rate of nutrient uptake. Application of 1-naphthylacetic acid （NAA） could partly replace the role of the removed apex, stimulated assimilate and nutrient deposition into the treated tissue, and enhanced the reduced plasma membrane ATPase activity of roots to the control level. However, treatment of the apex-removed plants with NAA could not rescue the reduced nutrient uptake rate and the shifted assimilates and nutrients partitioning caused by excision of the apex. Higher nutrient uptake rate of the intact plants could not be explained by root growth parameters, such as total root surface area and number of root tips. The results from the present study indicate that strong apex demand determined assimilates and nutrients partitioning and nutrient-uptake rate in tobacco （Nicotiana tabacum） plants.     

No evidence of a generalized potential ‘cost’ of apical dominance for species that have strong apical dominance

尚无证据表明顶端优势强的物种存在广义顶端优势潜在“成本”     

Manipulating resource allocation in plants

The distribution of nutrients and assimilates in different organs and tissues is in a constant state of flux throughout the growth and development of a plant. At key stages during the life cycle profound changes occur, and perhaps one of the most critical of these is during seed filling. By restricting the competition for reserves in Arabidopsis plants, the ability to manipulate seed size, seed weight, or seed content has been explored. Removal of secondary inflorescences and lateral branches resulted in a stimulation of elongation of the primary inflorescence and an increase in the distance between siliques. The pruning treatment also led to the development of longer and larger siliques that contained fewer, bigger seeds. This seems to be a consequence of a reduction in the number of ovules that develop and an increase in the fatty acid content of the seeds that mature. The data show that shoot architecture could have a substantial impact on the partitioning of reserves between vegetative and reproductive tissues and could be an important trait for selection in rapid phenotyping screens to optimize crop performance.     

Effects of resource availability, and fruit and ovule position on components of fecundity in Alliaria petiolata (Brassicaceae)

In four field and glasshouse experiments designed to alter the supply of resources through manipulation of nutrients, root tissue, leaf area and fruit number in Alliaria petiolata (Brassicaceae), more than 99% of ovules per plant showed signs of fertilization, suggesting that seed production in this plant was not pollen limited. However, in all treatments a significant proportion of fruits and seeds did not develop to maturity. Total fruit and seed production did not differ significantly from controls when plants were given nutrient supplements at flowering. Removal of 50–75% of the root tissue in 1-yr-old plants significantly reduced fruit set, but had no effect on individual seed development. Removal of cauline leaves significantly reduced most measures of fruit and seed production, suggesting that current photosynthate is critical for fruit and seed filling. Seed maturation was significantly affected by both fruit position within an infructescence and ovule position within a fruit. Basally located fruits and ovules (within fruits) developed more mature seeds than distally positioned fruits and ovules. Plants responded to removal of basal fruits by re-allocating resources to distal fruits that would normally have aborted. Our results suggest that fruits and seeds act as reproductive sinks competing for parental photosynthate. Patterns of resource allocation within infructescences and fruits were also modified by our experiments.     

Strigolactones are a new-defined class of plant hormones which inhibit shoot branching and mediate the interaction of plant-AM fungi and plant-parasitic weeds

Because plants are sessile organisms, the ability to adapt to a wide range of environmental conditions is critical for their survival. As a consequence, plants use hormones to regulate growth, mitigate biotic and abiotic stresses, and to communicate with other organisms. Many plant hormones function plei-otropically in vivo, and often work in tandem with other hormones that are chemically distinct. A newly-defined class of plant hormones, the strigolactones, cooperate with auxins and cytokinins to control shoot branching and the outgrowth of lateral buds. Strigolactones were originally identified as compounds that stimulated the germination of parasitic plant seeds, and were also demonstrated to induce hyphal branching in arbuscular mycorrhizal (AM) fungi. AM fungi form symbioses with higher plant roots and mainly facilitate the absorption of phosphate from the soil. Conforming to the classical definition of a plant hormone, strigolactones are produced in the roots and translocated to the shoots where they inhibit shoot outgrowth and branching. The biosynthesis of this class of compounds is regulated by soil nutrient availability, i.e. the plant will increase its production of strigolactones when the soil phosphate concentration is limited, and decrease production when phosphates are in ample supply. Strigolactones that affect plant shoot branching, AM fungal hyphal branching, and seed ger-mination in parasitic plants facilitate chemical synthesis of similar compounds to control these and other biological processes by exogenous application.     

Nicotine synthesis in Nicotiana tabacum L. induced by mechanical wounding is regulated by auxin

The effects of different kinds of mechanical wounding on nicotine production in tobacco plants were compared, with sand or hydroponics culture under controlled conditions. Both removal of the shoot apex and damage of the youngest unfolded leaves nos 1 and 2 by a comb-like brusher with 720 punctures caused an increase in nicotine concentration in whole plants at day 3, and reached its highest level at day 6. The nicotine concentration induced by excision of the shoot apex was much higher than that induced by leaf wounding. Both treatments also caused an increase in jasmonic acid (JA) concentration within 90 min in the shoot, followed by an increase in the roots (210 min), in which the JA concentration induced by leaf wounding was significantly higher than that induced by excision of the shoot apex. The increase in nicotine concentration occurred throughout the whole plant, especially in the shoot, while the increase in JA concentration in the shoot was restricted to the damaged tissues, and was not observed in the adjacent tissues. Removal of the lateral buds that emerged after excision of the shoot apex caused a further increase in nicotine concentrations in the plant tissues. Removal of mature leaves, however, did not cause any changes in nicotine concentration in the plant, even though the degree of wounding in this case was comparable with that occurring with apex removal. The results suggest that the nicotine production in tobacco plants was not correlated with the degree of wounding (cut-surface or punctures), but was highly dependent on the removal of apical meristems and hence on the major sources of auxin in the plant. Furthermore, immediate application of 1-naphthylacetic acid (NAA) on the cut surface after removing the shoot apex completely inhibited the increase both in nicotine in whole plants and in JA in the damaged stem segment and roots. Application of an auxin transport inhibitor around the stem directly under the shoot apex of intact plants also caused an increase in nicotine concentration in the whole plant. The results strongly suggest that auxin serves as a negative signal to regulate nicotine synthesis in roots of tobacco plants.     

Nicotine concentration in leaves of flue-cured tobacco plants as affected by removal of the shoot apex and lateral buds

It is believed that the nicotine concentration in tobacco is closely correlated with the amount of nitrogen (N) supplied.On the other hand,N uptake mainly occurs at the early growth stage,whereas nicotine concentration increases at the late growth stage,especially after removing the shoot apex.To identify the causes of the increased nicotine concentration in tobacco plants,and to compare the effects of different ways of mechanical wounding on nicotine concentration,field experiments were carried out in Fuzhou,Fujian Province in 2003 and 2004.Excision of the shoot apex had almost no influence on N content in the plant;however,it caused dramatic increases in nicotine concentration in leaves,especially in the middle and upper leaves.An additional increase of the nicotine concentration was obtained by removal of axillary buds.The wounding caused by routine leaf harvests,however,did not change the leaf nicotine concentration,and neither did reducing leaf harvest times.The present results revealed no direct relationship between N supply and nicotine concentration in tobacco leaves,and indicate that not all kinds of mechanical wounding were capable of stimulating nicotine synthesis in tobacco plants.Since nicotine production is highly dependent on the removal of apical meristems and hence on the major sources of auxin in the plant,and application of 1-naphthylacetic acid onto the cut surface of the stem after removing the shoot apex markedly decreased the nicotine concentration in different leaves and the total nicotine content in the plant,the results suggest that decreased auxin supply caused by removal of the shoot apex as a kind of mechanical wounding might regulate nicotine synthesis in the roots of tobacco plants.     

Shoot apex removal does not alter autoregulation of Modulation in soybean

The suppression of new nodule development in soybean (Glycine max (L.) Merr.) has been previously demonstrated to involve the shoot through reciprocal grafts between the wild-type cultivar Bragg and its supernodulating mutant nts382. Using the same grafting technique, but modified through the excision of the shoot apex region and emerging lateral shoots, we show here that autoregulation of nodule number still existed despite apex removal. This radical treatment lowered total nodule number per plant as well as root, shoot and nodule dry weight. Bragg shoots grafted onto nts382 roots gave wild-type nodulation (26 nodules, 15mg total nodule mass) as compared to nts382 shoots grafted onto Bragg roots (340 nodules, 277 mg total nodule mass). Specific nodule mass differed between supernodulating (about 0·5-1·0mg per nodule) and wild-type nodulating (2·3 mg per nodule) plants. In contrast to other growth characteristics, apex removal did not affect specific nodule size, except in plants with wild-type shoots and nts382 (supernodulation) roots. Apex removal only slightly affected the percentage of nodule weight per total root weight in nts382, but had a severe effect in wild type. Growth reductions varied between the normal and supernodulating plants. The fact that autoregulation of nodulation still functions in plants devoid of functional shoot apices suggests that the autoregulation signal may not be derived from the apex regions and that the leaf may be a likely source.     

Maternal phenotypic effects due to soil nutrient levels and sink removal in Arabidopsis thaliana (Brassicaceae)

Three levels of soil nutrients and systematic removal of racemes and siliques, intended to reduce reproductive sink size, induced maternal effects in a genetically uniform, inbred accession of Arabidopsis thaliana ecotype Col-0. Soil nutrient levels but not trimming treatments caused significant differences in maternal plant weight and number of seeds per silique. Trimming and increased soil nutrient level resulted in larger seeds. Germination rates were significantly affected only by the trimming main effect, while there were no significant effects on germination percentage. At 14 and 19 d harvest dates, soil nutrient level treatments resulted in significant differences in offspring aerial biomass, but this effect was nonsignificant at 24, 29, and 34 d harvest dates. The effect of trimming on offspring aerial biomass was significant at all harvest dates. No significant soil nutrient level × trimming interaction was observed on any harvest date. Phenotypic correlations among aerial biomass at each harvest date and seed weight of the six offspring classes were highly significant. Analyses of variance for germination and biomass data adjusted for seed weight showed no significant differences due to soil nutrient level, trimming, or their interaction, indicating that maternal effects influencing these variables were determined by seed weight. Relative growth rates were estimated from changes in biomass over time. The negative correlation between initial relative growth rate and rate of change of the relative growth rate was highly significant (R = -0.99; P < 0.01).     

Mycorrhizal infection of wild oats: maternal effects on offspring growth and reproduction

Summary The objective of this study was to determine whether infection of Avena fatua L. plants by the mycorrhizal fungus Glomus intraradices Schenck & Smith could influence the vigor of the offspring generation. Two experiments demonstrated that mycorrhizal infection of the maternal generation had slight but persistent positive effects on offspring leaf expansion in the early stages of growth. In two other experiments, mycorrhizal infection of mother plants had several long lasting effects on their offspring. Offspring produced by mycorrhizal mother plants had greater leaf areas, shoot and root nutrient contents and root:shoot ratios compared to those produced by non-mycorrhizal mother plants. Moreover, mycorrhizal infection of mother plants significantly reduced the weight of individual seeds produced by offspring plants while it increased the P concentrations of the seeds and the number of seeds per spikelet produced by offspring plants. The effects of mycorrhizal infections of maternal plants on the vigor and performance of offspring plants were associated with higher seed phosphorus contents but generally lighter seeds. The results suggest that mycorrhizal infection may influence plant fitness by increasing offspring vigor and offspring reproductive success in addition to previously reported increases in maternal fecundity.     

Effects of below- and above-ground herbivores on plant growth,flower visitation and seed set

Separate and combined effects of root and leaf herbivores on plant growth, flower visitation and seed set were tested in a factorial experiment using potted mustard, Sinapis arvensis, at an old fallow field. A 50% leaf removal by cabbageworms (Pieris rapae) when the seedlings had their first four leaves reduced plant height and shoot mass, and delayed the onset of flowering. Root herbivory by two wireworms (Agriotes sp.) over the whole experiment changed flower visitation; the number of flower visitors per plant was higher in plants with root herbivores than in plants without root herbivores. Combined leaf and root herbivory affected flowering period, number of fruits per plant and number of seeds per fruit. Plants attacked by leaf and root herbivores had a shorter flowering period and produced fewer fruits per plant than plants with root herbivores only. Although the experimental plants faced major herbivore-induced growth changes, plant reproduction (seed set and weight per plant) was similar in all treatments, documenting their ability to effectively compensate for leaf and root herbivory.     

DEVELOPMENTAL CHANGES IN ENERGY (ATP AND ADP) AND NUTRIENT LEVELS PRIOR TO SENESCENCE IN PODDED AND DEPODDED PEA PLANTS

Energy (ATP and ADP) levels in stem apices of depodded pea plants (Pisum sativum L. cv. Little Marvel) were significantly higher than those of podded plants during the pod-filling stage before whole plant senescence. This difference in energy content appeared before decreases in leaf chlorophyll and soluble proteins occurred in plants of both treatments. In contrast, the mineral nutrient levels (N, P, K, Ca, Mg, Fe, and Mn) in stem apices of plants from both treatments were similar. Energy levels in reproductive leaves from podded and depodded plants were similar. The mineral nutrients in leaves with fruits in their axils and similar leaves of depodded plants were comparable except for nitrogen. Plant growth measurements—dry weight, leaf area, leaf dry weight, root/shoot ratio—were significantly higher in depodded than podded plants. Whole plant senescence occurred significantly earlier in podded than in depodded pea plants.     

RESPONSE OF THE ANNUAL ABUTILON THEOPHRASTI MEDIC. (MALVACEAE) TO TIMING OF NUTRIENT AVAILABILITY

Because resource variability is likely to be high in disturbed environments, colonizing species should be opportunistic in their abilities to use resources at any time during the growing season. We examined the effects of timing of nutrient availability on seed quantity and quality in the colonizing annual Abutilon theophrasti. Plants were grown under six different nutrient treatments involving additions of different size, frequency and timing but having the same total amount of nutrients in all treatments. The plants showed an opportunistic response, being able to use nutrients supplied at different times for reproduction. However, when nutrient availability was delayed until fruit maturation had begun, seed number and total seed weight were reduced, production of mature fruits was delayed, and seed P concentrations were reduced. Seeds from the first two fruits of the plants had higher N concentration (mg/g), higher N content (mg/seed), lower P concentration, higher K content, and higher mean seed weight.     

VARIATION IN SEED WEIGHT AND ITS EFFECTS ON GERMINATION IN PASTINACA SATIVA L. (UMBELLIFERAE)

Pastinaca sativa (wild parsnip) produces seeds on the primary, secondary, and tertiary umbels of the flowering stalk. Within plants, variation in seed weight is about twofold. Secondary and tertiary seed weight is 73% and 50% of primary seed weight, respectively. Maximum variation in seed weight between plants is sixfold when tertiary seeds from a small plant are compared to primary seeds from a large plant. Within an umbel order, variation in seed weight between plants is correlated with plant size. Under autumn germinating conditions in the laboratory, final germination of seeds from different umbel orders does not differ but smaller seeds germinate more rapidly than larger seeds. Under spring germination conditions in the laboratory, significantly more primary and secondary seeds germinate than tertiary seeds and the rate of germination is independent of seed weight. Field germination of seeds from different umbel orders produces similar results except that in the spring both secondary and tertiary seed germination is lower than that of primary seeds. These results suggest that with respect to seed germination characteristics small seeds may have a competitive advantage over large seeds in the autumn because they germinate more quickly, but in the spring small seeds are at a disadvantage because they have lower overall germination. Because most germination in the field occurs in the spring, population recruitment from small seeds is likely to be substanially less than that from large seeds.     

Endozoochorous dispersal of aquatic plants: does seed gut passage affect plant performance?

The ingestion of seeds by vertebrates can affect the germinability and/or germination rate of seeds. It is, however, unclear if an earlier germination as a result of ingestion affects later plant performance. For sago pondweed, Potamogeton pectinatus, the effects of seed ingestion by ducks on both germinability and germination rate have been previously reported from laboratory experiments. We performed an experiment to determine the effects of seed ingestion by ducks on germination, seedling survival, plant growth and asexual multiplication. Both at the start and end of the winter, seeds were fed to three captive shovelers (Anas clypeata) and planted outdoors in water-filled containers. Plant biomass and its allocation to vegetative parts (shoot and roots), tubers, and seeds were determined in autumn. More duck-ingested seeds than control (uningested) seeds germinated in early winter, but this difference disappeared for seeds planted in late winter, when the treatments were first stratified for 3 mo. None of the variables for measuring seedling survival and plant performance varied between treatments. Under our experimental conditions (no herbivory or competition), ingestion by ducks in early winter resulted in increased performance for seeds surviving gut passage due to enhanced seed germinability, without other costs or benefits for the seedlings.     

Flooding effects on rapid responses of the invasive plant Alternanthera philoxeroides to defoliation

In experiments under controlled growth conditions it was examined how flooding affected the responses of the invasive plant Alternanthera philoxeroides to defoliation. In drained and flooded conditions, plants were subjected to five defoliation levels: 0, 10, 50, 90% removal of leaf tissue and apex removal (90% leaf tissue plus apical bud removal). Plants were harvested weekly for five weeks. In drained conditions, plant biomasses including total biomass, shoot biomass and root biomass after 50% defoliation rapidly recovered to the control plant level. They were significantly lower for the 90% defoliation and apex removal treatments compared to control plants throughout the experiment. In flooded conditions, total biomass and shoot biomass after 50% defoliation, 90% defoliation, and apex removal treatments could return to control plant levels before the end of the experiment. In 90% defoliation and apex removal treatments root to shoot biomass ratios of both drained and flooded plants were initially much higher than in control plants, but the difference disappeared rapidly. The final biomasses decreased with increased defoliation intensity in drained conditions, but no significant difference was generally found in any of the defoliation treatments in flooded conditions. The rapid re-growth of A. philoxeroides plants after defoliation may partly be responsible for its invasion success. However, defoliation capable of removing 90% of the leaf tissue may be desirable in restricting the growth of this invasive species in drained conditions.     

bushy, a dominant pea mutant characterised by short, thin stems, tiny leaves and a major reduction in apical dominance

The spontaneous, single-gene dominant, pea ( Pisum sativum L.) mutant bushy is characterised by short, thin stems, tiny leaves and a proliferation of basal lateral branches. We symbolised the dominant mutant allele bsh and the recessive wild-type allele BSH . Some effects were very large, e.g. the reduction in internode length was around 10-fold in pure mutant plants. The effect on branching was qualitative under our conditions as the wild-type did not branch and the mutant branched extensively. Analysis of epidermal cells indicated the reduction in internode length arose principally from a reduction in cell length. The bushy mutation also altered root morphology with a reduction in the number and length of lateral roots. Time to first open flower was increased but node of flower initiation was not affected. In a few cases, bushy plants died before producing an open flower even though tiny abortive flower buds were produced in the upper leaf axils. In pure mutant plants, individual seed weight was reduced by 30%, number of seeds per pod was reduced 3-fold, and seed number per plant was reduced 4-fold. However, pod size was essentially normal for a given seed content, and the flowers were fertile and of normal structure. Grafting studies showed the primary action of the bushy mutation occurred in the shoot. In summary, the reduced cell and shoot elongation, loss of apical dominance and a primary action in the shoot, all point toward auxin deficiency (or perceived deficiency) as a possible cause of the bushy phenotype. The overall characteristics of bushy make it a useful mutant for research on plant development.     

EFFECTS OF PARENTAL IDENTITIES AND ENVIRONMENT ON COMPONENTS OF CROSSING SUCCESS IN PHLOX DRUMMONDII

A controlled pollination study, involving a diallel crossing scheme among 15 Phlox drummondii individuals grown under either control, low water or low nutrient conditions, was performed. The results of this research demonstrate that, although plant size and flower number are responsive to the treatments, pollen germination percentage, the survival probability of developing seeds and the weight of mature seeds are buffered against water and nutrient deprivation. However, the identities of male and female parents did have significant effects on pollen germination and seed survival, and female identity had a significant effect on seed weight.     

Does attraction to frugivores or defense against pathogens shape fruit pulp composition?

Fruit traits evolve in response to an evolutionary triad between plants, seed dispersers, and antagonists that consume fruits but do not disperse seeds. The defense trade-off hypothesis predicts that the composition of nutrients and of secondary compounds in fruit pulp is shaped by a trade-off between defense against antagonists and attraction to seed dispersers. The removal rate model of this hypothesis predicts a negative relationship between nutrients and secondary compounds, whereas the toxin-titration model predicts a positive relationship. To test these alternative models, we evaluated whether the contents of nutrients and secondary compounds can be used to predict fruit removal by mutualists and pathogens in 14 bird-dispersed plants on a subtropical island in São Paulo state, southeastern Brazil. We selected eight to ten individuals of each species and prevented fruit removal by covering four branches with a net and left fruits on four other branches available to both, vertebrate fruit consumers and pathogens. The persistence of ripe fruits was drastically different among species for bagged and open fruits, and all fruit species persisted longer when protected against seed dispersers. We found that those fruits that are quickly removed by vertebrates are nutrient-rich, but although the attack rate of pathogens is also high, these fruits have low contents of quantitative defenses such as tannins and phenols. Thus, we suggest that the fruit removal rate by seed dispersers is the primary factor selecting the levels of fruit defense. Likewise, nutrient-poor fruits have low removal of seed dispersers and low probability of attack by pathogens. These species retain ripe fruits in an intact condition for a prolonged period because they are highly defended by secondary compounds, which reduce overall attractiveness. However, this strategy might be advantageous for plants that depend on rare or unreliable dispersers.