Competition and genetic background in a rapid-cycling cultivar of brassica rapa (Brassicaceae)

We examined the effects of a slight genetic disadvantage in a competitive situation by comparing the performance of inbred and outcrossed Brassica rapa plants over a range of neighbor densities, using a rapid-cycling, self-compatible cultivar of this species. We also examined the genetic correlations in performance among plants grown alone and with intra- and interspecific competitors. Competition had a strong effect on biomass and on the number of flowers produced, but differences in biomass between inbred and outcrossed plants were dependent on the competitive environment. Outcrossed plants outperformed those that were selfed only at intermediate densities of neighbors; at high densities and in the absence of competition inbred and outcrossed plants did not differ. For outcrossed families, performance without competitors and in intra- and interspecific competition were all highly correlated, but for selfed families, correlations were low, and performance in competition was not predictable from that of plants grown alone. Thus, the phenotypic expression of genetic differences may depend on the density of neighbors with which plants are grown.     

AN EXPERIMENTAL STUDY OF COMPETITIVE PERFORMANCE IN BRASSICA RAPA (CRUCIFERAE)

Growth chamber experiments with rapid-cycling Brassica rapa were designed to estimate the signs and magnitudes of the genetic correlations for plant performance in each of three conditions: no-competition (isolated plants), intraspecific competition, and interspecific competition with Raphanus sativa. Biomass and flower number were highest in the no-competition treatment, intermediate under intraspecific competition, and lowest under interspecific competition. Significant among-family variation in biomass and flower number was found under each regime. The mean family performance (biomass or flower number) in the no-competition treatment was significantly positively correlated with the performance in only one of the competitive treatments (for biomass in the intraspecific treatment). For both biomass and flower number there was a significant positive correlation between family means in the intra- and interspecific regimes. These correlations were greater in magnitude than those for the comparison between no-competition and competition (intra- or interspecific) treatments. Our results suggest that the importance of traits affecting plant performance is environment-dependent; the performance of a family grown without competition was a poor predictor of performance with competition, while the performance of families grown under intra- and interspecific competition was positively correlated.     

Sibling competition, size variation and frequency-dependent outcrossing advantage in Plantago coronopus

Many plants display limited seed dispersal, thereby creating an opportunity for sibling competition, i.e. fitness-determined interactions between related individuals. Here I investigated the consequences of intra-specific competition, by varying density and genetic composition of neighbors, on the performance of seedlings derived by selfing or outcrossing of the partially self-fertilizing plant Plantago coronopus (L.). Seedlings from eight plants, randomly selected from an area of about 50 m² in a natural population, were used in (i) a density series with either one, four or eight siblings of each cross type per pot and (ii) a replacement series with eight plants per pot where selfed and outcrossed siblings were grown intermixed in varying frequencies. Density had a pronounced effect on plant performance. But, except for singly grown individuals, no differences were detected between selfed and outcrossed progenies in vegetative and reproductive biomass. When grown intermixed, selfed offspring were always inferior to their outcrossed relatives. The magnitude of reduction in performance was dependent on the number of outcrossed relatives a selfed seedling had to compete with, giving rise to a frequency-dependent fitness advantage to outcrossed seedlings. The major result of this study is (i) that the relative fitness of inbred progeny is strongly affected by the type of competitors (inbred or outbred) and (ii) that inbreeding depression varies according to the density and frequency of outbred plants and could be considered as a density- and frequency-dependent phenomenon. It is argued that sibling competition, due to the small genetic neighborhood of P. coronopus, might be an important selective force in natural populations of this species.     

Does competition with wind‐pollinated species alter Echium plantagineum's attractiveness to a common pollinator Bombus terrestris?

1. In insect‐pollinated plants, pollinator attraction is influenced by flowers (e.g. number, size) and their associated rewards (e.g. pollen, nectar). These traits can depend on plant interactions. Indeed, below‐ground competition between plants can lead to a decrease in flower or reward production in insect‐pollinated species. 2. Wind‐pollinated plants, in particular, which are almost never studied in plant–pollinator networks, can alter insect‐pollinated plants' attractiveness through competition for nutrients. The response of pollinators to such changes has never been investigated. 3. A pot experiment was carried out in which an insect‐pollinated species, Echium plantagineum, was grown in binary mixture with three wind‐pollinated species selected to exert a panel of competitive interactions. Below‐ground competition was controlled using dividers limiting interspecific root competition. Floral traits of E. plantagineum (i.e. flower production, floral display size, flower size and nectar production) were measured. For each species mixture, the visits (i.e. first visit, number of visits, 10‐min sequences) of Bombus terrestris individuals released in a flight cage containing two pots were followed, one with and one without below‐ground competition. 4. Below‐ground competition significantly affected nectar's sucrose concentration but did not influence flower and nectar production. Likewise, pollinator visits were not influenced by below‐ground competition. Competitor identity significantly influenced flower and reward production of E. plantagineum, with a decrease in the presence of the most competitive wind‐pollinated species. A tendency for faster flower visitation events was also detected in the presence of the least competitive competitor. This study raises new questions regarding the influence of wind‐pollinated plants on plant–pollinator interactions.     

Ecophysiological Responses to Neighbor Removal in an Old-Field and a Prairie in Northeastern Kansas,USA

An ecophysiological approach was used to determine if competition can be detected among plants in a recently abandoned old-field and in a native tallgrass prairie in northeastern Kansas. In situ photosynthetic parameters and water potentials (Ψ) of target plants were measured 1-2 d after neighbor (intra- and interspecific) removal as well as 1-4 weeks later, and compared with values for plants with neighbors. Only two of the six study species (four old-field and two prairie species) responded to removal of neighboring plants, and only after several weeks had elapsed. Net photosynthetic rates (P_N) and stomatal conductances (g_s) of Ambrosia trifida in an old-field increased after removal of both intra- and interspecific neighbors. For Apocynum cannabinum, another old-field species, P_N of target plants without neighbors was significantly higher than that of target plants with neighbors. For both these species, values of Ψ were not different between target plants with and without neighbors, suggesting that increased availability of nutrients may have been responsible for the observed ecophysiological responses. Though numerous past studies indicate that competition is a major factor influencing plants in old-field and in prairie communities, the experimental approach used in this study revealed that neighbor removal had only limited effects on ecophysiology of the target plants in either community. This revised version was published online in June 2006 with corrections to the Cover Date.     

Aggregative Root Placement: A Feature During Interspecific Competition in Inland Sand-Dune Habitats

Segregation of roots is frequently observed in competing root systems. However, recently, intensified root growth in response to a neighbouring plant has been described in pot experiments [Gersani M, Brown J S, O'Brien E E, Maina G M and Abramsky Z 2001. J. Ecol. 89, 660–669]. This paper examines whether intense root growth towards a neighbour (aggregation) plays a role in competitive interactions between plant species from open nutrient-poor mid-European sand ecosystems. In a controlled field-competition experiment, root distribution patterns of intra- and interspecific pairs as well as single control plants of Corynephorus canescens, Festuca psammophila, Hieracium pilosella, Hypochoeris radicata and Conyza canadensis were investigated after one growing season. Under intraspecific competition plants tended to segregate their root systems, while under interspecific competition most species tended to aggregate roots towards their neighbours even at the expense of root development at the opposite competition-free side of the target. Preference of a root aggregation strategy over the occupation of competition-free soil in interspecific competition emphasizes the importance of contesting between individuals in relation to mere resource acquisition. It is suggested that in the presence of a competitor the plants might use root aggregation as a defensive reaction to maintain a strong competitive response and exclusive access to the resources of already occupied soil volumes.     

Pollination of Ipomopsis aggregata (Polemoniaceae): effects of intra- vs. interspecific competition

Although plants may simultaneously experience intra- and interspecific competition for pollination, their relative strength has rarely been experimentally evaluated. Yet because intra- and interspecific competition can be caused by different mechanisms, their effect on the ecology and evolution of plants may differ. To determine the relative strength of intra- and interspecific competition for pollination, I manipulated the presence of heterospecifics and density of conspecifics using Ipomopsis aggregata as the focal species. All plots contained I. aggregata and Castilleja linariaefolia, but C. linariaefolia inflorescences were removed from half of the plots to create the heterospecifics-absent treatment. Within each plot, all I. aggregata inflorescences were removed from a 5-m radius around a focal plant to create a low conspecific density experimental unit, and a group of 12 I. aggregata plants/1 m² was designated as a high conspecific density unit. Conspecific pollen deposition was reduced when C. linariaefolia was present but was not influenced by I. aggregata density. Although seed set per fruit was reduced by 17% when C. linariaefolia was present, it was not significantly influenced by either treatment. Interspecific competition for pollination is stronger than intraspecific competition in the I. aggregata–C. linariaefolia system, but neither process appears to influence plant fitness.     

Effect of arbuscular mycorrhiza on inter- and intraspecific competition of two grassland species

We were interested in the role of arbuscular mycorrhiza (AM) in the competition between plants of different sizes. A pot experiment of factorial design was established, in which AM root colonization and competition were used as treatments. Five-week-old Prunella vulgaris seedlings were chosen as target plants (i.e. plants whose response to competition was studied) and the following (13 replicates of each) were used as neighbours: (1) a large, 10-week-old P. vulgaris, (2) two P. vulgaris seedlings, and (3) a large, 10-week-old Fragaria vesca. In the experiment where small neighbours were grown together with small target plants, competition did not reduce target plant weight significantly, compared to the other two treatments. The competitive effects of large neighbours were significant, regardless of species (both older neighbours reduced the weights of target plants similarly), but there was a clear difference between intra- and interspecific competition when plants were mycorrhizal. In intraspecific competition with a large neighbour, the target plant shoot weight was reduced 24% when inoculated with AM. Thus, AM amplified rather than balanced intraspecific competition. In interspecific competition with old F. vesca, the shoot weights of target plants were 22% greater when inoculated with AM than when non-mycorrhizal. The results showed that, for given soil condition, AM might increase species diversity by increasing competitive intraspecific suppression and decreasing the interspecific suppression of small plants by larger neighbours.     

Genetic analysis of interspecific incompatibility in Brassica rapa

In interspecific pollination of Brassica rapa stigmas with Brassica oleracea pollen grains, pollen tubes cannot penetrate stigma tissues. This trait, called interspecific incompatibility, is similar to self-incompatibility in pollen tube behaviors of rejected pollen grains. Since some B. rapa lines have no interspecific incompatibility, genetic analysis of interspecific incompatibility was performed using two F₂ populations. Analysis with an F₂ population between an interspecific-incompatible line and a self-compatible cultivar ‘Yellow sarson’ having non-functional alleles of S-locus genes and MLPK, the stigmas of which are compatible with B. oleracea pollen grains, revealed no involvement of the S locus and MLPK in the difference of their interspecific incompatibility phenotypes. In QTL analysis of the strength of interspecific incompatibility, three peaks of LOD scores were found, but their LOD scores were as high as the threshold value, and the variance explained by each QTL was small. QTL analysis using another F₂ population derived from selected parents having the highest and lowest levels of interspecific incompatibility revealed five QTLs with high LOD scores, which did not correspond to those found in the former population. The QTL having the highest LOD score was found in linkage group A02. The effect of this QTL on interspecific incompatibility was confirmed by analyzing backcrossed progeny. Based on synteny of this QTL region with Arabidopsis thaliana chromosome 5, a possible candidate gene, which might be involved in interspecific incompatibility, is discussed.     

Increase of histidine content in Brassica rapa subsp. oleifera by over-expression of histidine-rich fusion proteins

An approach that enables the increase of the quantity of a specific amino acid in crop plants is reported. Oleosin gene from Arabidopsis thaliana or 30K movement protein gene of Tobacco mosaic virus (TMV; genus Tobamovirus) were cloned under the control of napin or hybrid promoters, and in fusion to synthetic poly-histidine (poly-His) sequences for transformation into spring turnip rape (Brassica rapa subsp. oleifera; synonym to B. campestris). The most stable expression cassettes for the poly-His production prior to the plant transformation were selected by analyzing the protein expression in in vitro translation and in transient plant expression systems using GFP as marker. Expression of the poly-His-constructs in transgenic Brassica rapa plants was analyzed using dot and western blotting and PCR. The constructs were stably expressed in the third generation of the transgenic plant lines. Histidine content was measured from the seeds of the transgenic plants, and some plant lines had more than 20% increase in histidine content compared to wild type. The methodology may be widely applicable to increase the content of any amino acid in crop plants including those encoded by rare codons.     

INBREEDING DEPRESSION IN HYDROPHYLLUM APPENDICULATUM: ROLE OF MATERNAL EFFECTS,CROWDING, AND PARENTAL MATING HISTORY

This paper examines several aspects of the expression of inbreeding depression in an outcrossing, obligately biennial plant, Hydrophyllum appendiculatum (Hydrophyllaceae). The amount of inbreeding depression detected was small during the first year of life but increased with age and had significant effects on adult size and reproductive traits. The lack of significant inbreeding depression during early growth is likely due to the overriding influence of maternal environmental effects on seed size and seedling growth. However, as maternal effects decreased with age, the seedling's own genotype became a more important determinant of its fate. To examine whether the expression of inbreeding depression was sensitive to ecological conditions, selfed and outcrossed seedlings were grown alone or with other H. appendiculatum seedlings. No inbreeding depression was detected in the plants grown alone. In contrast, under competitive conditions, outcrossed seedlings were significantly larger than selfed seedlings by the end of the first growing season. To address whether parental mating history influences the amount of inbreeding depression expressed, I examined the consequences of two successive generations of selfing on seed set and seed weight. The amount of inbreeding depression increased following the second generation of selfing. In the first generation, seed set and seed weight differed by less than 5% between selfed and outcrossed progeny. However, both traits were 15% greater for outcrossed plants after two generations. These results indicate that the alleles responsible for the reductions in these traits were not purged and suggest the action of multiple loci with deleterious effects.     

Additive transgene expression and genetic introgression in multiple green-fluorescent protein transgenic crop × weed hybrid generations

The level of transgene expression in crop × weed hybrids and the degree to which crop-specific genes are integrated into hybrid populations are important factors in assessing the potential ecological and agricultural risks of gene flow associated with genetic engineering. The average transgene zygosity and genetic structure of transgenic hybrid populations change with the progression of generations, and the green fluorescent protein (GFP) transgene is an ideal marker to quantify transgene expression in advancing populations. The homozygous T₁ single-locus insert GFP/Bacillus thuringiensis (Bt) transgenic canola (Brassica napus, cv Westar) with two copies of the transgene fluoresced twice as much as hemizygous individuals with only one copy of the transgene. These data indicate that the expression of the GFP gene was additive, and fluorescence could be used to determine zygosity status. Several hybrid generations (BC₁F₁, BC₂F₁) were produced by backcrossing various GFP/Bt transgenic canola (B. napus, cv Westar) and birdseed rape (Brassica rapa) hybrid generations onto B. rapa. Intercrossed generations (BC₂F₂ Bulk) were generated by crossing BC₂F₁ individuals in the presence of a pollinating insect (Musca domestica L.). The ploidy of plants in the BC₂F₂ Bulk hybrid generation was identical to the weedy parental species, B. rapa. AFLP analysis was used to quantify the degree of B. napus introgression into multiple backcross hybrid generations with B. rapa. The F₁ hybrid generations contained 95–97% of the B. napus-specific AFLP markers, and each successive backcross generation demonstrated a reduction of markers resulting in the 15–29% presence in the BC₂F₂ Bulk population. Average fluorescence of each successive hybrid generation was analyzed, and homozygous canola lines and hybrid populations that contained individuals homozygous for GFP (BC₂F₂ Bulk) demonstrated significantly higher fluorescence than hemizygous hybrid generations (F₁, BC₁F₁ and BC₂F₁). These data demonstrate that the formation of homozygous individuals within hybrid populations increases the average level of transgene expression as generations progress. This phenomenon must be considered in the development of risk-management strategies.Communicated by J. Dvorak     

Crossability of Brassica tournefortii and B. rapa, and morphology and cytology of their F1 hybrids

The crossability between Brassica tournefortii (TT, 2n = 20) and Brassica rapa (AA, 2n = 20) and the cytomorphology of their F₁ hybrids were studied. Hybrids between these two species were obtained only when B. tournefortii was involved as a female parent. The hybrid plants were intermediate for most of the morphological attributes and were found to be free from white rust under field conditions. The F₁ plants showed poor pollen fertility, although occasional seed set was achieved from open pollination. Self-pollination or backcrosses did not yield any seeds in these plants. The occurrence of chromosome association ranging from bivalents (0–7), trivalents (0–2) to a rare quadrivalent (0–1) in the dihaploid hybrids indicates pairing between the T and A genomes. The homoeologous pairing coupled with seed set in the F₁ plants offer an opportunity for interspecific gene transfers from B. tournefortii to B. rapa and vice-versa through interspecific hybridization. Received: 3 July 2000 / Accepted: 22 September 2000     

Inbreeding effects on mating system traits for two species of Lupinus (Leguminosae)

The consequences of inbreeding for reproductive traits were investigated for two closely related annual lupines that differ in their mating system. Lupinus bicolor (Leguminosae) is a primarily selling species while Lupinus nanus outcrosses at intermediate rates. A controlled crossing program was used for each species to produce selfed and outcrossed progeny. These progeny were then grown in a greenhouse and scored for the date of first flower, flower morphology, and autofertility. Selfed progeny of L. bicolor produced significantly smaller flowers but did not differ from outcrossed progeny for the remaining traits. Selfed progeny of L. nanus produced flowers that significantly differed in shape and had fewer ovules than the flowers of outcrossed progeny. Selfed progeny of L. nanus also had significantly lower rates of autofertility in comparison to outcrossed progeny. The significant effects of inbreeding on these mating system traits may indicate the presence of directional dominance at the loci underlying these characters. The consequences of these direct effects of inbreeding on reproductive traits for plants growing in natural populations may include nonadaptive changes in the outcrossing rate between generations.     

喜旱莲子草和接骨草竞争对模拟增温的响应北大核心CSCD

研究外来入侵植物与本地植物种竞争对气候变暖的响应,对于预测未来气候变化背景下入侵植物的入侵趋势、理解其入侵机制以及筛选生态替代种具有重要的意义。以入侵我国的外来植物喜旱莲子草(Alternanthera philoxeroides)和本地植物种接骨草(Sambucus chinensis)为材料,通过两种植物单栽、纯栽和混栽,采用红外辐射加热器模拟增温,研究了两种植物竞争对模拟增温的响应。结果表明:(1)在模拟增温期间(2013年5–12月),增温组空气平均温度比不增温组提高了0.47℃,相对湿度降低了1.87%;(2)混栽的喜旱莲子草除根冠比与单栽无显著差异外,其余各生物量和根系形态指标均显著低于单栽喜旱莲子草;无竞争、种间竞争和种内竞争三种竞争间,接骨草除根冠比、细根与总根生物量比、比根长和比根表面积无显著差异外,其余指标均呈现无竞争>种间竞争>种内竞争的趋势;(3)无竞争、种间竞争和种内竞争三种条件下,喜旱莲子草各指标在增温和不增温处理间差异均不显著,而接骨草总生物量和根生物量在无竞争和种间竞争条件下增温处理均显著低于不增温处理,在种内竞争条件下则相反;(4)增温使接骨草的相对拥挤系数降低,接骨草对温度升高反应敏感,而喜旱莲子草则表现出一定的适应性。由此推测,在中度遮阴陆生生境中,接骨草有望成为喜旱莲子草生物替代控制的材料。     

Responses of an oligophagous beetle species to rearing for several generations on alternative host‐plant species

1. Imprinting and rearing on alternative host‐plant species over several generations may affect performance and preferences of herbivorous insects. In this study, the adjustment potential to alternative hosts was investigated in the oligophagous beetle Phaedon cochleariae (F.) (Coleoptera: Chrysomelidae). 2. A population that had been reared for several generations on Brassica rapa was split and transferred to either one of the alternative Brassicaceae species, Sinapis alba and Nasturtium officinale, or was kept on B. rapa for continuous rearing. After 10 generations, a subset of the Sinapis and Nasturtium populations was tested again on the original host species, B. rapa. Larval performance and adult preferences were tested. 3. In the second generation, beetles performed better (higher body mass and faster larval development) on B. rapa than on the two alternative species. No evidence for imprinting was found. After 10 generations of selection on the alternative plant species, the performance of the beetles on these species improved. This effect was more pronounced on S. alba than on N. officinale. However, the performance on B. rapa after 10 generations of selection on the alternative species was unaffected, and was nearly equal to the insects kept continuously on B. rapa, with slight deviations only in the lines selected on N. officinale. 4. Most females preferred to oviposit on B. rapa independent of their experience, suggesting a genetic fixation and/or a positive relationship between larval performance and adult preference. Preferences may have been also driven by physical leaf characteristics.     

远缘杂交油菜核不育系的创建及其细胞学和形态学研究

在甘蓝型油菜与诸葛菜以及芥菜型油菜与诸葛菜属间杂交后代中分别发现1个和3个不育材料,经杂交和多代近交育成了相应的甘蓝型油菜不育系。通过对核不育系体细胞鉴定表明,所有新发现的不育系染色体数为38,均已恢复到甘蓝型油菜。这些不育系绝大部分花粉母细胞（PMC）在中期Ⅰ、后期Ⅰ和后期Ⅱ 3个时期染色体行为表现正常,但不同时期的PMC均会出现一定比例的异常现象,主要表现为染色体落后或染色体桥等。这些不育系属于单核败育型,不育株与可育株的花器形态差异明显,不育系还存在不同程度的死蕾等特点。通过对花器生长过程的研究,发现不育株雌蕊生长随雄蕊败育进程逐渐加快,而可育株雌蕊生长则存在两个生长缓慢阶段。此外,文章还讨论了这些不育系的应用前景。     

Genetic changes in a novel breeding population of Brassica napus synthesized from hundreds of crosses between B. rapa and B. carinata

Introgression of genomic variation between and within related crop species is a significant evolutionary approach for population differentiation, genome reorganization and trait improvement. Using the Illumina Infinium Brassica 60K SNP array, we investigated genomic changes in a panel of advanced generation new‐type Brassica napus breeding lines developed from hundreds of interspecific crosses between 122 Brassica rapa and 74 Brassica carinata accessions, and compared them with representative accessions of their three parental species. The new‐type B. napus population presented rich genetic diversity and abundant novel genomic alterations, consisting of introgressions from B. rapa and B. carinata, novel allelic combinations, reconstructed linkage disequilibrium patterns and haplotype blocks, and frequent deletions and duplications (nonrandomly distributed), particularly in the C subgenome. After a much shorter, but very intensive, selection history compared to traditional B. napus, a total of 15 genomic regions with strong selective sweeps and 112 genomic regions with putative signals of selective sweeps were identified. Some of these regions were associated with important agronomic traits that were selected for during the breeding process, while others were potentially associated with restoration of genome stability and fertility after interspecific hybridization. Our results demonstrate how a novel method for population‐based crop genetic improvement can lead to rapid adaptation, restoration of genome stability and positive responses to artificial selection.     

Expression of a Chinese cabbage cysteine proteinase inhibitor, BrCYS1, retards seed germination and plant growth in transgenic Tobacco plant

Phytocystatins are plant cysteine proteinase inhibitors that regulate endogenous and heterologous cysteine proteinases of the papain family. A cDNA encoding the phytocystatin BrCYS1 (Brassica rapa cysteine proteinase inhibitor 1 ) has been isolated from Chinese cabbage (B. rapa subsp.pekinensis) flower buds. In order to explore the role of this inhibitory enzyme, tobacco plants (Nicotiana tabacum L. cv. Samson) containing altered amounts of phytocystatin were generated by over-expressingBrCYS1 cDNA in either the sense or the antisense configuration. The resulting plants hadin vitro enzyme inhibitory activities that were over 10% of those detected in wild type plants. The transgenic plants exhibited retarded seed germination and seedling growth and a reduced seed yield, whereas these properties were enhanced in antisense plants. These data suggest that BrCYS1 participates in the control of seed germination, post-germination and plant growth by regulating cysteine peptidase activity.