Which adaptations of some invasive Ludwigia spp. (Rosidae,Onagraceae) populations occur in contrasting hydrological conditions in Western France?

Biological invasions in wetlands by Water Primroses (Ludwigia grandiflora ssp. hexapetala and L. peploides ssp. montevidensis) are an increasing problem especially in wet meadows. The aim of this paper is to quantify differences between species and the adaptation of these amphibious plants in order to establish the consequences for site managers. The hypotheses were: (i) that these species differ in biomass and morphology, and (ii) that terrestrial habitats determine particular adaptations such as reduced biomass, more roots and a bushy form. Biomasses from five sites were collected, following seasonal changes and biological traits were measured on selected plants. Comparing species in aquatic habitats, L. peploides produced less biomass than L. grandiflora. Comparing habitats for L. grandiflora, the biomass in meadows was almost twice that in dykes, particularly due to old stems, the converse of the hypothesis. Terrestrial forms were characterized by a bushy morphology, with shorter internodes and stems, as well as more secondary ramifications, while in aquatic habitats the plant architecture was simpler. Models were built linking morphology to species, sites and water conditions. Adaptations to terrestrial habitats of a particularly resistant form with a significant potential biomass are practical constraints for farmers and managers.     

Co-existence of three species of amphibious plants in relation to spatial and temporal variation: Investigation of plant responses

Plant responses in Ludwigia peploides (Kunth) Raven ssp. montevidensis (Spreng.) Raven and Marsilea mutica Mett., and the introduced Myriophyllum aquaticum (Vellozo) Verdc. were observed and related to co-existence of the species in a freshwater lagoon in the Sydney basin. Water levels vary in the lagoon, and plants of all three species grow readily from stem fragments on freely draining, waterlogged and submerged soils. In Marsilea, sporelings develop rapidly from ruptured sporocarps when wet. In Ludwigia, the seeds germinate in white and in red light, but not in darkness or far-red light. They germinate readily on soil and in water, floating or when held submerged. At 30°C, germination is rapid, at 40°C slower and less, and at 10°C nil, but seeds imbibed at 10°C and then exposed at 15°C have greater and more rapid germination than seeds maintained at 15°C. In relation to seasonal variation, plants of all three species grew at 10, 20, 30 and 40°C on freely draining, waterlogged and submerged soils, with the optimum temperature for growth apparently lower in Myriophyllum than in the other two species. In relation to water depth, plants of all three species grew when submerged to I m; the stems of Ludwigia and Myriophyllum elongating and reaching the water surface, and the petioles in Marsilea elongating bringing the leaf blades on to the surface, each within a week during summer. The floating leaf blades in Marsilea increase in size with depth of submergence of the subtending shoots. Mean relative growth rates (R?) under the most favourable experimental conditions were least in plants of Marsilea, the species occurring along more exposed parts of the shoreline in the lagoon. It was concluded that, while water levels in the lagoon were stable, differences in growth form between the species were probably more critical in their co-existence than quantitative differences in their growth: the floating leaves of Marsilea, being more resistant to wind and wave action than the floating shoots of Ludwigia and Myriophyllum, occupy the more exposed parts of the lagoon 'S edge. The most sheltered parts are occupied by the shoots of Myriophyllum which ride high on the water and overtop shoots of Ludwigia. Shoots of Ludwigia appear to be more resistant than those of Myriophyllum to removal and damage by wind and wave action, and achieve their greatest abundance on the water surface in situations of intermediate exposure to wind and wave action. Using Grubb's (1977) classification of niches, in Bushells Lagoon the three species appear to be primarily differentiated in habitat niche, less in regeneration niche and even less in phenological niche. In the lagoon, their habitat niches apparently overlap in water depth but are differentiated along a gradient of exposure.     

Co-existence of three species of amphibious plants in relation to spatial and temporal variation: Field evidence

Changes in vegetation of Bushells Lagoon, a freshwater body fed intermittently by the Hawkesbury River in the Sydney basin, were studied over three scales of time: broad fluctuations in areas occupied by emergent plants over 28 years; fluctuations in their percentage cover over 21 months; and, over 6 months, seasonal variation in biomass of emergent plants of two native species Ludwigia peploides (Kunth) Raven ssp. montevidensis (Spreng.) Raven and Marsilea mutica Mett., and the introduced Myriophyllum aquaticum (Vellozo) Verde. Differences in distribution and abundance of plants observed between the three species were related to environmental variation in time and space. Wind and wave action varied along the shoreline, and emergent shoots of Myriophyllum were most abundant in sheltered conditions, floating leaves of Marsilea most abundant in relatively exposed conditions and emergent shoots of Ludwigia most abundant in somewhat intermediate conditions in the shallows. Chemical characteristics of the water varied in time but, unlike water temperature, had no obvious seasonal pattern. Seasonal change occurred in the biomass of Marsilea, and to a lesser extent in that of Myriophyllum, with smaller proportions of leaf material present in winter than in summer. Though water levels were apparently stable over a period of some years (1965–74), at other times they varied greatly through flooding and drought with related major change in cover of the three species. Co-existence of the three species in the lagoon may be related to differences between their plants in establishment and growth following disturbance (regeneration niche), tolerance of wind and wave action (habitat niche), or seasonal growth (phenological niche).     

The breakdown of macrophytes in a reservoir wetland

Weight losses from leaf laminae and petioles of Nelumbo lutea (Wild.) Pers., and from leaves of Ludwigia leptocarpa (Nutt.) Hara. and Typha angustifolia L., were measured by exposing air-dried leaf material in nylon mesh bags at upper (exposed 25 days, inundated 40 days) and lower (inundated 154 days) wetland sites in a Texas reservoir. No significant differences (P < 0.05) were found between sites, so data were combined to yield estimates of plant litter breakdown for the entire wetland. Breakdown rates (in percentage of ash free dry weight lost per day) for the 4 litter types were: Nelumbo leaves — 0.0108 ± 0.0016; Ludwigia — 0.0050 ± 0.0007; Typha — 0.0047 ± 0.0006; Nelumbo — petioles 0.003 ± 0.0010. Times required for a 95% loss of litter, based on an exponential model, are 278 days, 600 days, 638 days and 909 days for these 4 litter types, respectively. These rates are comparable to those reported for emergent aquatic macrophytes in other lakes and wetlands. Colonization of decaying wetland plants was dominated by the gatherers, Chironominii (54%) and Caenis sp. (26%).     

Temporal variations of physicochemical and microbiological parameters in three freshwater ecosystems (southeastern France) invaded by Ludwigia spp

In France, two amphibious hydrophytes of alien Ludwigia (Onagraceae) have for about the past twenty years been causing serious ecological and economic problems: L. peploides (Kunth) Raven et L. grandiflora (Michaux) Greuter & Burdet. This bacteriological and physicochemical study, focused on three different Mediterranean aquatic ecosystems, reveals, for the first time, a direct negative impact of these American invaders. During summer, while plant growth is intensive, and the appearance in the water column of anoxic conditions and production of toxic compounds may be observed, notably in L. grandiflora stands. The toxicity is linked to a proliferation of sulphate-reducing bacteria producing sulphides that are very harmful for aquatic organisms.     

Ludalbin,a new lactone from Artemisia ludoviciana

Artemisia ludoviciana Nutt. ssp. albula (Woot.) Keck, a member of the vulgaris complex, contains a sesquiterpene lactone, ludalbin, which is 8-α-acetoxydouglanine.     

Introgressive hybridization between Plantago major L. taxa

Occurrence and direction of introgressive hybridization between Plantago major taxa were tested. Plantago major plants were collected from 10 Egyptian locations. Four populations of two European taxa were used for comparison. These are ecologically and geographically separated and were identified as the subspecies Plantago major ssp. major and Plantago major ssp. intermedia. In the Egyptian populations, most individuals fall within the variation range of P. major ssp. intermedia. Only one population, from Burg El-Arab, morphologically resembled clearly P. major ssp. major and showed some ISSR fragments that characterize pure populations of this taxon. All individuals of population 2 (Alexandria) and some individuals of populations 1, 3, 5, 6 and 10 (Alexandria, Aswan) morphologically corresponded to P. major ssp. intermedia. All individuals collected from Egypt had ISSR fragments characterizing both pure P. major ssp. major and pure P. major ssp. intermedia. Most of these individuals had a higher percentage of intermedia-type fragments than major-type fragments, suggesting that P. major ssp. intermedia in Egypt shows some introgression towards P. major ssp. major. The pure populations were distinct from each other, while the Egyptian populations were intermediate according to principal coordinate analysis (PCoA) of ISSR data. In the populations collected from Egypt, major and intermedia cannot be seen as separate species. The study suggests that the dominant taxa introgressed to the minority population(s). Taxon frequency may be a key component in determining the direction of introgression.     

Lactobacillus delbrueckii ssp. lactis and ssp. bulgaricus: a chronicle of evolution in action

Background

Lactobacillus delbrueckii ssp. lactis and ssp. bulgaricus are lactic acid producing bacteria that are largely used in dairy industries, notably in cheese-making and yogurt production. An earlier in-depth study of the first completely sequenced ssp. bulgaricus genome revealed the characteristics of a genome in an active phase of rapid evolution, in what appears to be an adaptation to the milk environment. Here we examine for the first time if the same conclusions apply to the ssp. lactis, and discuss intra- and inter-subspecies genomic diversity in the context of evolutionary adaptation.

Results

Both L. delbrueckii ssp. show the signs of reductive evolution through the elimination of superfluous genes, thereby limiting their carbohydrate metabolic capacities and amino acid biosynthesis potential. In the ssp. lactis this reductive evolution has gone less far than in the ssp. bulgaricus. Consequently, the ssp. lactis retained more extended carbohydrate metabolizing capabilities than the ssp. bulgaricus but, due to high intra-subspecies diversity, very few carbohydrate substrates, if any, allow a reliable distinction of the two ssp. We further show that one of the most important traits, lactose fermentation, of one of the economically most important dairy bacteria, L. delbruecki ssp. bulgaricus, relies on horizontally acquired rather than deep ancestral genes. In this sense this bacterium may thus be regarded as a natural GMO avant la lettre.

Conclusions

The dairy lactic acid producing bacteria L. delbrueckii ssp. lactis and ssp. bulgaricus appear to represent different points on the same evolutionary track of adaptation to the milk environment through the loss of superfluous functions and the acquisition of functions that allow an optimized utilization of milk resources, where the ssp. bulgaricus has progressed further away from the common ancestor.

Electronic supplementary material

The online version of this article (doi:10.1186/1471-2164-15-407) contains supplementary material, which is available to authorized users.     

Reprint of: Saprolegnia strains isolated from river insects and amphipods are broad spectrum pathogens

Saprolegnia species are destructive pathogens to many aquatic organisms and are found in most parts of the world. Reports based on phylogenetic analysis suggest that Saprolegnia strains isolated from aquatic animals such as crustaceans and frogs are close to Saprolegnia strains isolated from infected fish or fish eggs and vice versa. However, it has often been assumed that host specificity occurs for each individual isolate or strain. Here we demonstrate that Saprolegnia spp. can have multiple hosts and are thus capable of infecting different aquatic organisms. Saprolegnia delica, Saprolegnia hypogyna, and 2 strains of Saprolegnia diclina were isolated from aquatic insects and amphipods while S. delica, Saprolegnia ferax, Pythium pachycaule, and a Pythium sp. were isolated from the water of a medium to fast flowing river. The ITS region of the rRNA gene was sequenced for all isolates. In challenge experiments, all four isolates from insects were found to be highly pathogenic to eggs of Atlantic salmon (Salmo salar) and embryos of the African clawed frog (Xenopus laevis). We found that Saprolegnia spp. isolated from salmon eggs were also able to successfully establish infection in nymphs of stonefly (Perla bipunctata) and embryos of X. laevis). These results suggest that Saprolegnia spp. are capable of infecting multiple hosts, which may give them an advantage during seasonal variation in their natural environments.     

Attraction of the potential biocontrol agent Altica cyanea by volatile compounds of three species of Ludwigia weeds from rice fields

Ludwigia adscendens (L.) Hara, Ludwigia parviflora Roxb., and Ludwigia octovalvis (Jacq.) Raven (Onagraceae) are abundant weeds in rice fields in India. These weeds compete with rice for resources in fields and this results in reduction of grain yield. Altica cyanea (Weber) (Coleoptera: Chrysomelidae) is a biocontrol agent of the three rice-field weeds. Hence, it is relevant to study host preference of A. cyanea using volatile cues of these three weeds. Therefore, we attempted to identify volatiles from leaves of the three Ludwigia species attracting A. cyanea, which could be used as an attractant during early emergence of the weeds in rice fields. In Y-tube olfactometer assays, A. cyanea females were more attracted to natural volatiles of plants after 48 h of feeding by adults than to volatiles of undamaged plants. The volatile organic compounds from undamaged plants, and plants after 6 and 48 h of feeding by A. cyanea were identified and quantified by gas chromatography and mass spectrometry (GC–MS) and GC-flame ionization detection (FID), respectively. In total, 25, 29, and 29 volatile compounds were detected in headspaces of undamaged L. adscendens, L. parviflora, and L. octovalvis, respectively, whereas 32, 35, and 34 compounds, respectively, were detected after 48 h of feeding by A. cyanea. Methyl jasmonate predominated among the volatile compounds in all treatments, but this compound was not attractive to A. cyanea. Females were attracted by synthetic blends of 3-hexanol, α-pinene, linalool oxide, and phytol in amounts mimicking those in each of the three Ludwigia species after 48 h of feeding by A. cyanea. The blends mimicking L. adscendens and L. parviflora included geraniol, whereas the blend mimicking L. parviflora also included 1-tridecanol. These synthetic blends may be helpful to monitor A. cyanea in biocontrol programmes.     

Abiotic influences on dissolved oxygen in a riverine environment

Dissolved oxygen (DO) is an important parameter in riverine health. Periphyton and/or macrophytes are frequently the drivers behind the fluctuation of DO levels in aquatic environments; however, the effects of abiotic and biotic factors on biomass and in turn on DO may be variable from river to river. The objective of this paper is to understand which factors govern DO levels in terms of daily minimum DO (DO_min) and daily DO variation (ΔDO) in a major wastewater-impacted river using statistical data analysis and modeling. Both climatic conditions (reflected in water temperature) and hydrometric conditions (flow) were major factors influencing DO_min and ΔDO. The effect of flow on DO_min and ΔDO was discontinuous, depending on the flow magnitude. Nutrient loading from wastewater effluent was not identified to have a significant impact on a daily basis; however, their role over large time scales is unclear. In the data-driven modeling approaches, the non-linear approach using multiple-layer perceptron neural network, which has very flexible architecture, was superior to the linear approach used (multiple linear regression). Although nutrients are likely related to the DO_min and ΔDO, flow and temperature were sufficient to obtain robust prediction in the data-driven modeling. This approach is useful to model and understand complicated processes when their governing mechanisms are not well presented in conceptual- or physically based models.     

Temporal and nonlinear dispersal patterns of <Emphasis Type="Italic">Ludwigia hexapetala</Emphasis> in a regulated river

Rivers are vulnerable to biological invasion due to hydrologic connectivity, which facilitates post-entry movement of aquatic plant propagules by water currents. Ecological and watershed factors may influence spatial and temporal dispersal patterns. Field-based data on dispersal could improve risk assessment models and management responses. Ludwigia hexapetala, an invasive emergent macrophyte, provides a case study for understanding dispersal patterns throughout a watershed. The species spreads via hydrochory and is increasingly imposing detrimental ecological and economic impacts within watersheds of the United States and Europe. We investigated morphology of shoot fragments and their dispersal in the Russian River watershed of California, capturing shoot fragments of L. hexapetala during repeated summer surveys at five locations in the river and quantifying their morphological traits that predict establishment success. Highly variable capture counts suggest the importance of pulse disturbance events in local dispersal of L. hexapetala. Unexpectedly, dispersing propagule pressure was nonlinear, with more shoot fragments captured in the middle rather than lower river. Captured fragments in the middle river were twice the length of fragments captured in the lower river and bore 83% more stem nodes, characteristics associated with greater establishment success. Our results support development of spatially targeted management, outreach, and prevention efforts that could lead to decreased propagule pressure in the watershed.     

Sulfated flavonoids and leaf morphology of the Halophila ovalis—H. minor complex (hydrocharitaceae) in the Pacific Islands and Australia

Sulfated flavonoids showed a systematic pattern among Halophila populations of the Pacific and Australia. Large-leaved plants, typical of H. ovalis (R.Br.) Hook. f., in the Palau Islands (Belau) and in Australia contained sulfated flavonoids, but small-leaved plants that have received various treatments under H. ovalis, including ssp. bullosa (Setchell) den Hartog and ssp. hawaiiana (Doty & Stone) den Hartog, and under H. minor (Zoll.) den Hartog or H. ovata Gaud. in American Samoa, Western Samoa, Tonga, Fiji, Wallis I., New Caledonia, Yap, Guam, Hawaii and Australia lacked sulfated flavonoids. Plants collected in their natural habitats as well as those grown in experimental cultures showed the characteristic patterns of these secondary compounds. Although the large-leaved plants have the potential for producing the largest leaves in the complex, they are not separable from small-leaved plants in leaf dimensions or in number of pairs of cross veins under all environmental conditions. The production of sulfated flavonoids and the potential for greater leaf length are the characteristics that can be used to separate typical H. ovalis and ssp. australis (Doty & Stone) den Hartog from other members of the complex.     

Relationships between the biomass production of invasive <Emphasis Type="Italic">Ludwigia</Emphasis> species and physical properties of habitats in France

Ludwigia grandiflora ssp. hexapetala and L. peploides ssp. montevidensis have invaded many types of fresh water ecosystem in France leading to serious problems of management of these macrophytes. The prediction of extracted plant biomass quantities for every control method could considerably help in plant removal and selecting management choices. The paper presents results of the research program “Biological Invasions” granted by the French Environmental Agency concerning the Ludwigia species (2003–2006): “Characterization of the relations between biotopes, plant populations and human activities: Implications for management”. The objective of this work is: (1) to compare environmental variables in the western part of France where colonised water bodies are widespread, (2) to link these parameters with the evaluation of standing crops and (3) to propose implications for management planning and control methods. Measured dry biomass of Ludwigia spp. varied from 200 g DM m⁻² in shallow lakes to 4,500 g DM m⁻² in a meander of a eutrophicated river. Results depend on the scale of the analysis: within sites, between-sites, at different seasons or years. In sunny conditions, optimal for these plants and in nutrient rich areas, biomass is high. In general, exposure to wind, waves and currents reduces the standing crop. For plant production as well as removal work, the status of plant development and water level, are key factors.     

Utilisation des lichens dans la mise en évidence des éléments traces présents dans les cours d'eau

The authors demonstrated the possibility to use lichens as biodetectors of the presence of copper and cadmium in rivers. The authors showed that the lichen biomass fixes rapidly metals in solution. This caption is proportional to the quantity of metal present, without any toxic threshold being observed, even at high concentration levels. Furthermore, Dermatocarpon luridum (With.) Laundon syn. D. weberi (Ach.) Mann was a better biodetector of metallic micropolluants than other aquatic macrophytes (brypophytes and phanerogames). Transplantations of this lichen were made in the Vienne river (Massif Central, France) in the upstream and downstream section of an electrolytic copper refining plant. The results revealed an aquatic pollution generated by this industry.     

Two Distinct Plastid Genome Configurations and Unprecedented Intraspecies Length Variation in the accD Coding Region in Medicago truncatula

We fully sequenced four and partially sequenced six additional plastid genomes of the model legume Medicago truncatula. Three accessions, Jemalong 2HA, Borung and Paraggio, belong to ssp. truncatula, and R108 to ssp. tricycla. We report here that the R108 ptDNA has a ∼45-kb inversion compared with the ptDNA in ssp. truncatula, mediated by a short, imperfect repeat. DNA gel blot analyses of seven additional ssp. tricycla accessions detected only one of the two alternative genome arrangements, represented by three and four accessions each. Furthermore, we found a variable number of repeats in the essential accD and ycf1 coding regions. The repeats within accD are recombinationally active, yielding variable-length insertions and deletions in the central part of the coding region. The length of ACCD was distinct in each of the 10 sequenced ecotypes, ranging between 650 and 796 amino acids. The repeats in the ycf1 coding region are also recombinationally active, yielding short indels in 10 regions of the reading frames. Thus, the plastid genome variability we report here could be linked to repeat-mediated genome rearrangements. However, the rate of recombination was sufficiently low, so that no heterogeneity of ptDNA could be observed in populations maintained by single-seed descent.     

Allelopathic potential of two invasive alien Ludwigia spp.

The allelopathic potential of two invasive alien Ludwigia [Onagraceae: L. peploides (Kunth) Raven and L. grandiflora (Michaux) Greuter and Burdet], that have developed quasi-monotypic stands in many aquatic ecosystems in France, was investigated. Since allelopathy involves the release of compounds into the environment, the water of monospecific experimental cultures was directly tested against two target species: Lactuca sativa L., the standard cultivar for bioassays, and Nasturtium officinale R. Brown, a resistant and widespread native hydrophyte. The treatment was carried out at the three main phases of development of both Ludwigia in February, May and August. For each experiment, the germination, mortality and culture yield percentages, the seedling growth (radicle and hypocotyl elongation) and the health of 15-day-old-seedlings were measured. The water of each Ludwigia tank induced: (1) a decrease in germination for watercress in August (control: 68.6%, L. peploides: 48.6%, L. grandiflora: 61.1%); (2) an increase in mortality in May only for watercress (control: 3.4%, L. peploides: 13.5%, L. grandiflora: 12%) and in August for both target species (up to 22.3% vs. 3% for lettuce and 27% vs. 12.5% for watercress); (3) a disturbance of seedling elongation for lettuce in all seasons; and (4) a seedling chlorosis of both target species, particularly in May and August. This study showed that L. peploides and L. grandiflora possess an allelopathic activity that influences the water quality throughout the year. Combined with the various competitive attributes, allelopathy may contribute to the great success of these two invasive Ludwigia in Europe. In threatened wetland communities of the Mediterranean area, in particular, allelopathy might have an important impact by diminishing the seedling survival of the most vulnerable species.     

The genetics of domestication of yardlong bean, Vigna unguiculata (L.) Walp. ssp. unguiculata cv.-gr. sesquipedalis

Background and Aims

The genetics of domestication of yardlong bean [Vigna unguiculata (L.) Walp. ssp. unguiculata cv.-gr. sesquipedalis] is of particular interest because the genome of this legume has experienced divergent domestication. Initially, cowpea was domesticated from wild cowpea in Africa; in Asia a vegetable form of cowpea, yardlong bean, subsequently evolved from cowpea. Information on the genetics of domestication-related traits would be useful for yardlong bean and cowpea breeding programmes, as well as comparative genome study among members of the genus Vigna. The objectives of this study were to identify quantitative trait loci (QTLs) for domestication-related traits in yardlong bean and compare them with previously reported QTLs in closely related Vigna.

Methods

Two linkage maps were developed from BC₁F₁ and F₂ populations from the cross between yardlong bean (V. unguiculata ssp. unguiculata cv.-gr. sesquipedalis) accession JP81610 and wild cowpea (V. unguiculata ssp. unguiculata var. spontanea) accession TVnu457. Using these linkage maps, QTLs for 24 domestication-related traits were analysed and mapped. QTLs were detected for traits related to seed, pod, stem and leaf.

Key Results

Most traits were controlled by between one and 11 QTLs. QTLs for domestication-related traits show co-location on several narrow genomic regions on almost all linkage groups (LGs), but especially on LGs 3, 7, 8 and 11. Major QTLs for sizes of seed, pod, stem and leaf were principally located on LG7. Pleiotropy or close linkage of genes for the traits is suggested in these chromosome regions.

Conclusions

This is the first report of QTLs for domestication-related traits in yardlong bean. The results provide a foundation for marker-assisted selection of domestication-related QTLs in yardlong bean and enhance understanding of domestication in the genus Vigna.