Timing of seed germination and the reproductive effort in Xanthium canadense

The effect of different dates of germination on the timing of flowering and the final reproductive yield was examined in a short-day annual plant Xanthium canadense (cocklebur). Delays in germination of 30 and 60 days deferred flower initiation by 2 and 9 days, respectively. Although plants that germinated later were smaller because of the shorter growing period, the reproductive yields did not show as much reduction as the vegetative biomass. The reproductive effort (RE, defined as the ratio of final reproductive yield to the vegetative biomass at the end of the growing season) increased 1.5 and 2.5 times with delays in germination of 30 and 60 days, respectively. A simple model of plant growth was used to analyse the factors involved in the control of RE, which depends only on the dry mass productivity and its partitioning in the reproductive phase, and is independent of the productivity and partitioning in the vegetative phase. Since relative allocation of dry mass to the reproductive part in the reproductive phase was similar for plants with different germination dates, the different REs could be ascribed mainly to differences in productivity of the vegetative parts in the reproductive period. The dependence of RE on plant size is discussed.     

Patterns of resource allocation in the dioecious alpine herb Aciphylla simplicifolia (Apiaceae)

Abstract Patterns of reproductive and vegetative biomass allocation were compared in male and female plants of the alpine herb Aciphylla simplicifolia. Male and female plants had similar vegetative biomass but differed in the pattern of resource allocation. Inflorescences of males and females were similar in weight at the time of flowering, but differed in biomass allocation to some structures within the inflorescences, particularly those associated with ovule production and pollinator attraction (number and size of flowers). At the time of fruit production, female inflorescences were 2.6 times heavier than at flowering with developing fruit six times heavier than flowers. In addition to the increase in biomass allocated to structures associated with the provisioning and dissemination of seed, support structures (main and side stalks) were also heavier. As a result of this additional investment of resources at the time of fruit production, the reproductive effort (RE) of female plants was much higher than that of males: 37% of above ground biomass compared with 21% for males. Differences in RE did not change with plant size; however, allocation to reproduction appeared to be a constant proportion of biomass over nearly all plant sizes sampled. These results show that sex‐specific resource allocation can be a complex of temporal and morphological patterns.     

Genetic interactions in the control of flowering time and reproductive structure development in tomato (Solanum lycopersicum)

Different tomato (Solanum lycopersicum) mutants, affected in flowering time, reproductive structure or plant architecture, were crossed to produce double mutants in order to investigate gene interactions in flowering regulation in this autonomous species with a sympodial growth habit. The compound inflorescence: uniflora, uniflora: self pruning, uniflora: blind, and jointless: uniflora double mutants all produced solitary flowers like their uniflora parent, instead of inflorescences. All double mutants were late flowering. uniflora: blind and uniflora: self pruning had flowering times intermediate between those of their two parents. jointless: uniflora and compound inflorescence: uniflora flowered later than uniflora, the mutant with the most delayed flowering. All double mutants developed strong lateral shoots at node levels approximately corresponding to the level at which their parent cultivars initiated their first reproductive structure, which is a typical trait of uniflora. These results suggest that the UNIFLORA gene acts upstream of the other investigated genes in controlling flowering in tomato, and that floral transition of the primary shoot and floral transition of sympodial segments are regulated differently.     

Multiplex CRISPR/Cas9-mediated mutagenesis of alfalfa FLOWERING LOCUS Ta1 (MsFTa1) leads to delayed flowering time with improved forage biomass yield and quality

Alfalfa (Medicago sativa L.) is a perennial flowering plant in the legume family that is widely cultivated as a forage crop for its high yield, forage quality and related agricultural and economic benefits. Alfalfa is a photoperiod sensitive long-day (LD) plant that can accomplish its vegetative and reproductive phases in a short period of time. However, rapid flowering can compromise forage biomass yield and quality. Here, we attempted to delay flowering in alfalfa using multiplex CRISPR/Cas9-mediated mutagenesis of FLOWERING LOCUS Ta1 (MsFTa1), a key floral integrator and activator gene. Four guide RNAs (gRNAs) were designed and clustered in a polycistronic tRNA–gRNA system and introduced into alfalfa by Agrobacterium-mediated transformation. Ninety-six putative mutant lines were identified by gene sequencing and characterized for delayed flowering time and related desirable agronomic traits. Phenotype assessment of flowering time under LD conditions identified 22 independent mutant lines with delayed flowering compared to the control. Six independent Msfta1 lines containing mutations in all four copies of MsFTa1 accumulated significantly higher forage biomass yield, with increases of up to 78% in fresh weight and 76% in dry weight compared to controls. Depending on the harvesting schemes, many of these lines also had reduced lignin, acid detergent fibre (ADF) and neutral detergent fibre (NDF) content and significantly higher crude protein (CP) and mineral contents compared to control plants, especially in the stems. These CRISPR/Cas9-edited Msfta1 mutants could be introduced in alfalfa breeding programmes to generate elite transgene-free alfalfa cultivars with improved forage biomass yield and quality.     

Patterns of reproductive effort in male and female shrubs of Oemleria cerasiformis: a 6-year study

1 We monitored flowering and fruiting of individual male and female plants of Oemleria cerasiformis over a 6-year period in a population in western Canada, and calculated fruit set (percentage of pistils maturing) and reproductive effort (RE) (gram of reproductive tissue per gram of leaf).
2 Over 6 years, male O. cerasiformis had on average much lower total RE, but much higher RE at flowering, than females.
3 In males, strong correlations between RE and light suggested that investment in reproduction was largely determined by light levels. There were strong positive correlations of RE between years, with no evidence of periodic fluctuations.
4 In females, in contrast to males, RE at flowering was not related to light. However, fruit set was strongly correlated with light. Flowering RE and fruit set were uncorrelated in females, indicating that these are affected by different factors.
5 Correlations of RE between years in females, although often significant, were lower than in males, indicating that RE fluctuates more between years in females than in males and may respond to past levels of RE. Flowering may reflect adjustments in response to past reproduction, or may be controlled by resources other than light. Fruit set was not significantly related to previous RE.
6 The greater total RE of females and their limited ability to adjust fruit set are probably major factors contributing to the greater mortality rates of females and the male-biased sex ratios in O. cerasiformis .     

Growth and reproduction of Arabidopsis thaliana in relation to storage of starch and nitrate in the wild-type and in starch-deficient and nitrate-uptake-deficient mutants

We have investigated the interactions between resource assimilation and storage in rosette leaves, and their impact on the growth and reproduction of the annual species Arabidopsis thaliana. The resource balance was experimentally perturbed by changing (i) the external nutrition, by varying the nitrogen supply; (ii) the assimilation and reallocation of resources from rosette leaves to reproductive organs, by cutting or covering rosette leaves at the time of early flower bud formation, and (iii) the internal carbon and nitrogen balance of the plants, by using isogenic mutants either lacking starch formation (PGM mutant) or with reduced nitrate uptake (NU mutant). When plants were grown on high nitrogen, they had higher concentrations of carbohydrates and nitrate in their leaves during the rosette phase than during flowering. However, these storage pools did not significantly contribute to the bulk flow of resources to seeds. The pool size of stored resources in rosette leaves at the onset of seed filling was very low compared to the total amount of carbon and nitrogen needed for seed formation. Instead, the rosette leaves had an important function in the continued assimilation of resources during seed ripening, as shown by the low seed yield of plants whose leaves were covered or cut off. When a key resource became limiting, such as nitrogen in the NU mutants and in plants grown on a low nitrogen supply, stored resources in the rosette leaves (e.g. nitrogen) were remobilized, and made a larger contribution to seed biomass. A change in nutrition resulted in a complete reversal of the plant response: plants shifted from high to low nutrition exhibited a seed yield similar to that of plants grown continuously on a low nitrogen supply, and vice versa. This demonstrates that resource assimilation during the reproductive phase determines seed production. The PGM mutant had a reduced growth rate and a smaller biomass during the rosette phase as a result of changes in respiration caused by a high turnover of soluble sugars ( Caspar et al. 1986 ; W. Schulze et al. 1991 ). During flowering, however, the vegetative growth rate in the PGM mutant increased, and exceeded that of the wild-type. By the end of the flowering stage, the biomass of the PGM mutant did not differ from that of the wild-type. However, in contrast to the wild-type, the PGM mutant maintained a high vegetative growth rate during seed formation, but had a low rate of seed production. These differences in allocation in the PGM mutant result in a significantly lower seed yield in the starchless mutants. This indicates that starch formation is not only an important factor during growth in the rosette phase, but is also important for whole plant allocation during seed formation. The NU mutant resembled the wild-type grown on a low nitrogen supply, except that it unexpectedly showed symptoms of carbohydrate shortage as well as nitrogen deficiency. In all genotypes and treatments, there was a striking correlation between the concentrations of nitrate and organic nitrogen and shoot growth on the one hand, and sucrose concentration and root growth on the other. In addition, nitrate reductase activity (NRA) was correlated with the total carbohydrate concentration: low carbohydrate levels in starchless mutants led to low NRA even at high nitrate supply. Thus the concentrations of stored carbohydrates and nitrate are directly or indirectly involved in regulating allocation.     

Modelling reproductive effort in sub-and maritime Antarctic mosses

A comparison is made of the reproductive effort (RE), considered as the investment in sporophyte relative to gametophyte biomass, of eight species of moss occurring at sub-and maritime Antarctic sites. Six of the species showed smaller sporophytes and game-tophytes at the climatically more extreme maritime Antaretic sites and one species showed no size difference between regions. The remaining species, although showing no regional difference, showed some evidence of a reverse pattern, with higher altitude samples having greater biomass than lower altitude samples. Spore counts indicated a measure of compensation in maritime Antarctic samples, with no significant decrease in spore output in several species despite smaller sporophyte biomass. The relationship between sporophyte (S) and gametophyte (G) biomass within samples was described by an allometric curve (S=aG ^b ) which gave a better fit than a straight line for six species. This form of model allows comparisons of patterns of RE to be made between samples with non-or partially overlapping size distributions, even when the relationship involves size-dependence. An allometric curve was not appropriate for describing samples of one species (Andreaea regularis), and insufficient data were available to identify any relationship in Polytrichum alpinum. The exponent (b) differed between species, but there were no statistically significant differences between exponents from samples of the same species. Samples of two species could further be described by the same coefficient (a), indicating that they lie on the same curve. However, samples of three species from sub-Antarctic South Georgia gave significantly higher coefficients, indicating increased RE relative to maritime Antarctic populations.     

VARIATION IN SEX ALLOCATION AND FLORAL MORPHOLOGY IN IPOMOPSIS AGGREGATA (POLEMONIACEAE)

Intrapopulational variation in biomass allocation to male vs. female function was quantified for the hermaphroditic plant Ipomopsis aggregata in terms applicable to sex allocation models. The proportions of flower biomass put into the corolla and calyx averaged 0.59 and 0.20 and were relatively constant across plants. The proportions in the stamens and pistil averaged 0.13 and 0.08, with considerable variation among plants. Phenotypic gender at the time of flowering ranged from 0.34 to 0.77 female. Pistil dry weight was correlated with stigma exsertion. Stamen weight was correlated with corolla width, which influences male pollination success, and was also correlated with anther position and pollen production. Female reproductive success as estimated by seeds per flower showed no detectable relationship with initial allocation of biomass at the time of flowering, but decreased in accelerating fashion with the proportion of final biomass including seeds that was allocated to male function.     

Pleiotropic effects of flowering time genes in the annual crucifer Arabidopsis thaliana (Brassicaceae)

Variation in flowering time of Arabidopsis thaliana was studied in an experiment with mutant lines. The pleiotropic effects of flowering time genes on morphology and reproductive yield were assessed under three levels of nutrient supply. At all nutrient levels flowering time and number of rosette leaves at flowering varied among mutant lines. The relationship between these two traits depended strongly on nutrient supply. A lower nutrient supply first led to an extension of the vegetative phase, while the mean number of leaves at flowering was hardly affected. A further reduction resulted in no further extension of the vegetative phase and, on average, plants started flowering with a lower leaf number. At low nutrients, early flowering affected the timing of production of siliques rather than the total output, whereas late flowering was favorable at high nutrients. This may explain the fact that many plant species flower at a relatively small size under poor conditions. Flowering time genes had pleiotropic effects on the leaf length, number of rosette and cauline leaves, and number of axillary flowering shoots of the main inflorescence. Silique production was positively correlated with the number of axillary shoots of the main inflorescence; the number of axillary primordia appeared to have a large impact on reproductive yield.     

Age-specific residual reproductive value and reproductive effort in the Iceland scallop,Chlamys islandica (O.F. Müller)

Summary The possibility for a trade-off between age-specific residual reproductive value (RRV) and reproductive effort (RE) was investigated. Different ways of estimating RE did not affect conclusions which were: a) there is no clear trade-off between RRV and RE until the scallop has reached an age of about 13 years, i.e. after a reproductive life of about eight years; b) the larva of C. islandica is more important as a means of dispersal than as a means of maintaining the parent population.     

唐古特雪莲花部特征及生殖分配的海拔差异

为探究高山植物生殖分配策略以及分析唐古特雪莲花部特征对海拔梯度的响应机制,该研究利用采样调查法和烘干称重法,对分布在青藏高原东缘不同海拔14个居群的唐古特雪莲花部特征和生殖分配进行了研究。结果表明:(1)繁殖分配随个体大小(地上生物量和株高)的增大呈线性递减趋势。(2)花期植株地上生物量、株高、管状小花数目、繁殖器官及营养器官生物量均随海拔升高呈线性递减趋势,管状小花生物量随海拔升高呈线性递增趋势。(3)花期管状小花数量及大小、繁殖器官生物量与营养器官生物量、雄蕊重量和雌蕊重量以及花粉数目与花丝长度之间均存在权衡现象。由此推论:(1)海拔作为外界因子对唐古特雪莲花部特征具有显著的影响,个体大小对其繁殖分配也存在潜在调控作用。(2)在不同海拔梯度上,唐古特雪莲有效地整合了有限的资源,其适应性特征之一就是通过减小个体大小来削弱营养生长以达到促进生殖生长的目的。     

Breeding of Brassica rapa for Biogas Production: Heterosis and Combining Ability of Biomass Yield

The use of plant biomass as substrate for biogas production has recently become of major interest in Europe. Winter Brassica rapa produces high early biomass and could be grown as a pre-crop harvested early in the year followed by a second crop such as maize. The objectives of this study were to estimate heterosis and combining ability of 15 European winter B. rapa cultivars for biomass yield at end of flowering. A half-diallel without reciprocals was conducted among cultivars to produce 105 crosses. These crosses and their parents were evaluated in two years at two locations in Northern Germany. Data collected were days to flowering (DTF), fresh biomass yield (FBY), dry matter content (DMC) and dry biomass yield (DBY). The mean DBY was 5.3 t/ha for the parental cultivars and 5.6 t/ha for their crosses. The crosses surpassed on average their parents by 7.6% for FBY and 5.9% for DBY whereas DMC was 1.4% higher in the parents. Maximum mid parent heterosis was 21.0% for FBY and 30.4% for DBY. Analysis of variance showed that genetic variance was mainly due to specific combining ability (SCA). The correlation between parental performance and general combining ability (GCA) was 0.42** for FBY and 0.53** for DBY. In conclusion, the amount of heterosis in crosses between European winter B. rapa cultivars is not very high on average, but can be up to 30% in the best crosses. Selection of parental combinations with high specific combining ability to produce synthetic cultivars can rapidly improve biomass yield.     

Effects of shift in flowering time on the reproductive output of Xanthium canadense in a seasonal environment

We studied the effects of a change in flowering date on the reproductive output of a short-day annual plant, Xanthium canadense. The flowering date was changed by photoperiodic manipulation to 1 month earlier or later than the natural flowering date. Plants with the natural flowering date attained the highest reproductive output. For those flowering 1 month earlier or later, the reproductive output was decreased by 42% or 23%, respectively. The reproductive output was analyzed as the product of the biomass production during the reproductive period and its allocation to the reproductive organs. Although delay in flowering increased biomass production, it decreased its fractional allocation to the reproductive organs. The highest reproductive output in the natural flowering plants resulted from a compromise between these two effects of flowering. Plants flowering earlier had higher translocation rates to the reproductive organs and accelerated plant senescence. Later flowering caused a reduction in biomass translocation to the reproductive organs and thus extended the reproductive period. These experimental results are discussed in relation to the cost of reproduction and the optimal time for flowering that maximizes the final reproductive output. It is suggested that the natural flowering time maximized the reproductive output while minimizing the cost of reproduction. Received: 11 September 1997 / Accepted: 12 December 1997     

RESOURCE ALLOCATION AND REPRODUCTION IN POPULATIONS OF AMPHICARPUM PURSHII (GRAMINEAE)

Five New Jersey populations of Amphicarpum purshii (an annual panicoid grass) were investigated as to total allocation of biomass to reproduction and its distribution between aerial chasmogamous and subterranean cleistogamous inflorescences. Germination of both types of seed was documented, for the first time, in not only the field but under three laboratory temperature regimes. Overall, approximately 29 % of the shoot biomass was allocated to reproduction. On a population basis, subterranean inflorescences accounted for 37–100 % of the reproductive biomass, and these seed (5 × heavier but fewer in number) were the source of most surviving seedlings. Subterranean spikelets contained caryopses significantly more frequently than aerial spikelets. The ratio of the number of viable aerial seed to the number of viable subterranean seed increased from 0/4 to 4/2 with recentness and/or frequency of disturbance. As a pioneer species in secondary succession, Amphicarpum purshii produces a larger subterranean propagule, with greater seedling vigor and a higher probability of local reproductive success, and also a smaller aerial propagule in larger numbers with potentially greater genetic variability.     

Plant size,flowering synchrony and edge effects: What,how and where they affect the reproductive success of a Neotropical tree species

Plant reproductive success is supposedly influenced by phenology and individual size, which may be modified under edge effects. We tested if reproductive success, estimated by fruit set, in Senefeldera verticillata (Euphorbiaceae) is related to flowering synchrony and tree size, including plant height and circumference at breast height. The study was carried out in the interior and in edges of clearings for gas pipelines and electric lines of a lowland rainforest in south‐eastern Brazil. Monthly observations were performed during one reproductive season, of 19 individuals that grew at edges of electric lines and gas pipelines and at forest interior. Reproductive success was significantly higher at forest interior than at gas pipeline area; there was no significant difference between gas pipeline and electric line areas or between forest interior and electric line area. In the forest edges, only plant height was positively related to plant reproductive success. This is probably related to crown exposure to sunlight, which enhances flower production. At forest interior, reproductive success was positively influenced by the synchrony of flowering activity among neighbouring individuals. In contrast, flowering synchrony based on phenophase intensity negatively impacted reproductive success. Senefeldera verticillata shows temporal dioecy and is mainly pollinated by small social bees, and the high degree of flowering synchrony at low intensity may increase the number of mating partners and therefore enhance its reproductive success. Inside the forest fragments, individuals with thicker trunks showed lower reproductive success, which may be related to a loss of reproductive capacity of older individuals. Our results evidenced the complexity of responses experienced by tropical plants subjected to forest fragmentation because of linear clearings.     

Soil Patches of Inorganic Nitrogen in Subtropical Brazilian Plant Communities with Araucaria angustifolia

One-year old tubers of two hybrid calla lily (calla) cultivars (Zantedeschia ‘Pot of Gold’ and ‘Majestic Red’) were inoculated with the arbuscular mycorrhizal fungus (AMF), Glomus intraradices, or not, and grown at three different rates of phosphorus (P) supply to asses the effects of AMF-inoculation on plant development (time of shoot emergence and flowering), flowering (number, length and rate of flowering), and tuber biomass and composition over two growing cycles (2002, 2003). Tubers and flowers of calla responded differently to AMF inoculation. Differences in mycorrhizal responsiveness between cultivars was related to differences in P requirements for flower and tuber production, and the influence of P supply on resource allocation to different reproductive strategies. Inoculation increased shoot production and promoted early flowering, particularly in 2003. Inoculated plants also produced larger tubers than non-inoculated plants, but only increased the number of flowers per plant in 2003. High P supply also increased tuber biomass, but decreased the number of flowers per plant in 2002. Plants grown at a moderate P-rate, produced the most flowers in 2003. For ‘Majestic Red’, benefits from AMF were primarily in terms of tuber yield and composition, and AMF effects on marketable flower production could potentially have negative impact on production strategies for growers. Inoculation of ‘Pot of Gold’ primarily influenced flower production and aspects of tuber quality that caused detectable enhancement of tuber yield and flowering in the second growing cycle following inoculation (2003). The results of this study show that the responses of calla to AMF are partially a function of how nutrient supply alters resource allocation to sexual and vegetative reproduction. Whether AMF-induced changes in resource allocation to flowering and tubers significantly alters commercial productivity and quality of calla depends on the crop production goals (e.g. tubers, cut flowers or potted plants). The U.S. Government’s right to retain a non-exclusive, royalty free licence in and to any copyright is acknowledged.     

Identification of quantitative trait loci influencing aerial morphogenesis in the model legume <Emphasis Type="Italic">Medicago truncatula</Emphasis>

In many legume crops, especially in forage legumes, aerial morphogenesis defined as growth and development of plant organs, is an essential trait as it determines plant and seed biomass as well as forage quality (protein concentration, dry matter digestibility). Medicago truncatula is a model species for legume crops. A set of 29 accessions of M. truncatula was evaluated for aerial morphogenetic traits. A recombinant inbred lines (RILs) mapping population was used for analysing quantitative variation in aerial morphogenetic traits and QTL detection. Genes described to be involved in aerial morphogenetic traits in other species were mapped to analyse co-location between QTLs and genes. A large variation was found for flowering date, morphology and dynamics of branch elongation among the 29 accessions and within the RILs population. Flowering date was negatively correlated to main stem and branch length. QTLs were detected for all traits, and each QTL explained from 5.2 to 59.2% of the phenotypic variation. A QTL explaining a large part of genetic variation for flowering date and branch growth was found on chromosome 7. The other chromosomes were also involved in the variation detected in several traits. Mapping of candidate genes indicates a co-location between a homologue of Constans gene or a flowering locus T (FT) gene and the QTL of flowering date on chromosome 7. Other candidate genes for several QTLs are described. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Variation in Photosynthetic Rates and Biomass Productivity among Four Mulberry Cultivars

Among four mulberry (Morus alba L.) cultivars (K-2, MR-2, BC2-59, and S-13), highest net photosynthetic rate (P _N) was observed in BC2-59 while the lowest rates were recorded with K-2. Significant differences among the four cultivars were found in leaf area, biomass production, activities of ribulose-1,5-bisphosphate carboxylase and sucrose phosphate synthase, and glucose and sucrose contents. The P _N and the activities of photosynthetic enzymes in the four cultivars were significantly correlated with the growth and biomass production measured as leaf yield, total shoot mass, and aerial plant biomass.     

Differences in response to defoliation between males and females of Silene dioica

Summary The response by male and female plants to herbivory was studied by experimental defoliation of the dioecious perennial herb Silene dioica in a green-house. Male and female plants were defoliated prior to and during the early flowering phase at two intensities (50% and 100% of leaf-area removed) in two consecutive years. Defoliation resulted in a decrease in the number of flowers initiated in both sexes, while a larger delay of peak flowering and a higher mortality was observed in males compared to females. In female plants, severe defoliation resulted in a reduction in seed number per capsule and in seed size compared to control. Females showed a negative correlation between the production of flowers in the first and second season in all treatments, while flowering in males the first season was not correlated with flowering in the second season. Females also showed a lower frequency of flowering than males during the two seasons studied. However, during the flowering period, males allocated significantly more biomass to flowers than did females. This outcome supports the idea that females may have a higher total reproductive expenditure than males, but males have a higher reproductive effort during flowering. Male rosette leaves were significantly preferred by the generalist herbivore Arianta arbustorum in experiments. This preference was most pronounced in trials with leaves from fertilized plants compared to nonfertilized plants. A greater storage of resources in aboveground leaves during winter by males compared to females may explain the higher preference for male leaves and the higher male mortality following early defoliation. Furthermore, males are smaller than females and may have a lower ability than females to replace lost resources needed for reproduction when defoliated early in the season.