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Although the impacts of single-species invasions on resident communities have been widely studied, the ecological effects of co-invasion (simultaneous invasion by multiple alien species) remain largely unexplored. In this study, using an analytical framework, we investigate the individual and combined effects of two global plant invaders (Leucanthemum vulgare and Lupinus polyphyllus) on the aboveground vegetation (plant richness, diversity and biomass) and belowground soil variables (pH, salinity, electrical conductivity, total dissolved solids, organic carbon [OC], phosphorous, and potassium) of grassland communities in the Himalaya. We studied these ecological variables in comparable plots under the following four scenarios: both invaders absent (i.e. uninvaded), either of the two invaders present (i.e. single species invasion), and both invaders present (i.e., co-invasion). Our results show invasion effects vary with the invasion scenarios and the studied variables. The single species invasion plots had lower mean plant richness and diversity but higher mean soil OC than the uninvaded plots. The co-invasion plots had significantly increased mean plant biomass and soil OC, but had a non-significant effect on plant richness and diversity compared to the uninvaded plots. The overall combined effect of species' co-invasion was additive in nature, but the effect varied significantly with choice of the response variable. We found synergistic effects of co-invasion on species richness and diversity, antagonistic effects on soil pH, and additive effects on the remaining response variables when compared with the predicted additive effect of the two invaders. Our findings suggest that further spread of L. polyphyllus will impact the resident plant community more severely when co-invading with L. vulgare than when invading alone, thereby lending support to the invasional meltdown hypothesis. Nevertheless, antagonistic and additive impacts for soil pH and other response variables respectively hint towards other alternate mechanisms at play. Overall, our study showcases a proof-of-concept to empirically estimate the additive, antagonistic and synergistic impacts of plant co-invasions, and also provides a model for designing co-invasion studies on other taxa/systems/scales elsewhere. Finally, based on the insights gained from this study, we highlight priority areas of future research in species' co-invasions. [ABSTRACT FROM AUTHOR]

Drought stress is a common abiotic stress, and Lupinus polyphyllus presents strong adaptability, but its drought resistance mechanism has not been explored. This study used PEG-6000 to simulate drought stress, and the H2O2 content, O2− generation rate and MDA content were determined. Transcriptome sequencing and untargeted metabolome analyses were also carried out on an Iceland germplasm and American B germplasm under different drought stress durations. The results showed that the gene regulation range in the American B germplasm was greater, whether genes were upregulated or downregulated. And the number of genes in the American B germplasm was higher than that in the Iceland germplasm. Additionally, the Iceland germplasm produced less peroxide under PEG stress than the Iceland germplasm. The Iceland germplasm was more stable than the American B germplasm under PEG stress, which can be shown in two aspects: peroxide content and gene regulation quantity. Joint transcriptomics and metabolomics analysis showed that genes and metabolites related to secondary and carbon metabolism were mainly involved in the response of Lupinus polyphyllus to PEG-simulated drought stress. The metabolites mainly included phenylalanine, tyrosine, trans-2-hydroxycinnamate, starch synthase, 1,4-alpha glucan branching enzyme and glycogen phosphorylase, and genes mainly included COMT, F5H, REF1, CAD, UGT72E and TPS. These results provided genetic resources and a theoretical basis for further molecular breeding of Lupinus polyphyllus. [ABSTRACT FROM AUTHOR]

Drought stress is a common abiotic stress, and Lupinus polyphyllus presents strong adaptability, but its drought resistance mechanism has not been explored. This study used PEG-6000 to simulate drought stress, and the H2O2 content, O2− generation rate and MDA content were determined. Transcriptome sequencing and untargeted metabolome analyses were also carried out on an Iceland germplasm and American B germplasm under different drought stress durations. The results showed that the gene regulation range in the American B germplasm was greater, whether genes were upregulated or downregulated. And the number of genes in the American B germplasm was higher than that in the Iceland germplasm. Additionally, the Iceland germplasm produced less peroxide under PEG stress than the Iceland germplasm. The Iceland germplasm was more stable than the American B germplasm under PEG stress, which can be shown in two aspects: peroxide content and gene regulation quantity. Joint transcriptomics and metabolomics analysis showed that genes and metabolites related to secondary and carbon metabolism were mainly involved in the response of Lupinus polyphyllus to PEG-simulated drought stress. The metabolites mainly included phenylalanine, tyrosine, trans-2-hydroxycinnamate, starch synthase, 1,4-alpha glucan branching enzyme and glycogen phosphorylase, and genes mainly included COMT, F5H, REF1, CAD, UGT72E and TPS. These results provided genetic resources and a theoretical basis for further molecular breeding of Lupinus polyphyllus.

The article presents experimental data on the species composition of fungal and viral pathogens in the secondary range of Lupinus polyphyllus Lindl. Tobacco mosaic virus, bean yellow mosaic virus, bean common mosaic virus, and pea enation mosaic virus were diagnosed for the first time on L. polyphyllus. Issues related to the peculiarities of the adaptability of viruses to invasive plant species are discussed. The preventive role of virus vectors (Aphididae) in the expansion of viruses and the expansion of the spectrum of host plants (susceptible species) is emphasized. Interaction with vectors, including the nonspecific species, is one of the mechanisms for the adaptability of viruses, their expansion into new regions, and the formation of new pathosystems with invasive plant species. It was concluded that it is possible to predict the search for the most effective variants of harmful organisms on the basis of the analysis of the trophic relationships of vectors for the biocontrol of L. polyphyllus. [ABSTRACT FROM AUTHOR]

Lupines (Lupinus spp.) are a common plant species on western U.S. rangelands with several lupine species containing alkaloids that can be toxic and/or teratogenic to livestock. In North America, more than 150 lupine species are recognized with some ranches or grazing allotments containing multiple species. One or more of these lupine species may contain alkaloids that are teratogenic to cattle. Previous work has shown that lupine alkaloids can be detected in earwax of cattle grazing lupine infested rangelands. Our hypothesis is that earwax can be used to determine if cattle have been exposed to teratogenic alkaloids from multiple lupine species. Two lupine species, L. sericeus and L. polyphyllus , were present on a rangeland in east-central Idaho. The teratogen, anagyrine, was detected in L. sericeus and the teratogen, ammodendrine, was detected in L. polyphyllus plants collected on this rangeland. In this study, earwax was collected from 69 pregnant cows that had previously grazed a rangeland containing two different lupine species containing alkaloids that cause crooked calf syndrome (CCS). Anagyrine was detected in the earwax of all 69 cows sampled. Ammodendrine, was detected in the earwax of 28 of the 69 cows sampled. Earwax is a good non-invasive sample to aid in the diagnosis of cattle that have consumed lupine and does appear, in this case, to be a good diagnostic tool to differentiate between more than one lupine species that may be the cause of CCS. Concentrations of anagyrine or ammodendrine did not correlate with the incidence of CCS. [Display omitted] • Some lupines contain teratogenic alkaloids that cause crooked calf syndrome in cattle. • Earwax is a non-invasive specimen that can be used to detect alkaloids in cattle consuming lupine. • The teratogen, anagyrine, was detected in all earwax samples from cattle that grazed pastures containing Lupinus sericeus. • The teratogen, ammodendrine, was detected in earwax of 28 of 69 cows that grazed pastures containing Lupinus polyphyllus. • Earwax can be used to differentiate between more than one lupine species that may be the cause of crooked calf syndrome. [ABSTRACT FROM AUTHOR]

Glyphosate is the most widely used non‐selective herbicide in the world. Glyphosate residues in soil can affect plant quality by modifying plant physiology, hormonal pathways and traits, with potential consequences for plants' interactions with herbivores.We explored these indirect effects in the context of plant–herbivore interactions in a perennial, nitrogen‐fixing herb. We quantified leaf herbivory for glyphosate‐exposed and control plants grown in phosphorus‐fertilized and non‐fertilized soils, and assessed the impacts of glyphosate treatment on traits related to plant resistance against herbivores (leaf trichome density, leaf mass per area) and performance (aboveground biomass, root:shoot ratio, nodule number, nodule activity). Moreover, we conducted a laboratory feeding experiment to compare the palatability of leaves from glyphosate‐exposed and control plants to a generalist mollusc herbivore.Herbivore damage and intensity in situ increased during the growing season regardless of glyphosate or phosphorus treatment. Glyphosate treatment reduced leaf trichome density but had no effect on the other plant traits considered. Herbivore damage was negatively associated with leaf trichome density. The feeding experiment revealed no difference in the feeding probability of mollusc herbivores between glyphosate‐exposed and control plants. However, there was an interaction between glyphosate treatment and initial leaf area for leaf consumption by herbivores: leaf consumption increased with increasing leaf area in both groups, but at a lower rate for glyphosate‐exposed plants than for control plants.Our results show that glyphosate residues in soil have the potential to indirectly affect aboveground herbivores through changes in leaf quality, which may have mixed consequences for folivore damage. [ABSTRACT FROM AUTHOR]

A weed plant of the species Lupinus polyphyllus was found in settlements located on the territory of the natural park "Vepssky forest" of the Leningrad region. The plant is known as a transforming species in the southern regions of Russia. The distribution of L. polyphyllus within the Vepsian forest occurs not due to the migration of the species to the north along transport routes, as one might assume, but due to flight from gardens and the formation of spontaneous invasive populations. The goal of the study was to identify the molecular and biochemical characteristics of garden lupine at the northern borders of its secondary range. To interpret the obtained data, the task of the study also included a comparison of intraspecific variability between the "old" invasive populations (in Finland and Central Russia, studied by us earlier) and the "new" naturalizing population of L. polyphyllus in the Vepsian forest. The search for L. polyphyllus localities in the territory of the Vepssky Les natural park was carried out by the route method with geobotanical descriptions of experimental sites (5 m × 5 m). DNA was isolated from eight herbarium specimens of L. polyphyllus (MHA, LE) and fifteen specimens collected in the territory of the natural park "Vepssky Les". To reveal the internal structure and phylogenetic relationships in lupine populations, networks of nuclear and chloroplast haplotypes and cluster analysis (UPGMA) with the SplitsTree program were used. The total content of polyphenols and flavonoids in the leaves was determined spectrophotometrically. The low inter-locality variability of ITS indicates that the populations of L. polyphyllus in Central Russia and in the North of Russia (St. Petersburg and Vepsskaya Pushcha) have the same origin. Analysis of the chloroplast intergenic spacer (rpl32–trnL) indicates intrapopulation diversity and suggests the presence of microevolutionary processes near the northern limits of the secondary distribution range of L. polyphyllus. The high content of polyphenols and flavonoids in the leaves reveals the adaptive capabilities of lupine in the studied area. Evidence suggests that a neglected species may soon become invasive, as has already happened in other regions. [ABSTRACT FROM AUTHOR]

Invasive plant species pose significant ecological threats worldwide, affecting the stability and biodiversity of local ecosystems. As a result of their control, a considerable amount of plant biomass is produced, which can be used to produce various value-added products. Five different composts were prepared from three invasive plant species found in Latvia - Reynoutria japonica, Solidago canadensis, Lupinus polyphyllus. The stages of composting have been investigated and recommendations for process optimization have been made based on the quality characterization of the final compost. The quality of the prepared invasive plant biomass composts has been evaluated based on the main plant nutrient concentration, humic substance concentration, and mineral contents. The allelopathic lupin alkaloid concentration throughout the composting process has been evaluated and shows a consistent reduction. Obtained compost quality complies with the EU regulations for fertilizing products and soil amendments thus it can be considered equivalent to industrially produced compost and vermicompost. Seed germination tests confirm that compost prepared from invasive plants is suitable for plant growth and comparable to commercial composts. Based on pilot-scale composting results, recommendations for invasive plant composting have been suggested., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Plant phenology, i. e. the timing of life cycle events, is related to individual fitness and species distribution ranges. Temperature is one of the most important drivers of plant phenology together with day length. The adaptation of their phenology may be important for the success of invasive plant species. The present study aims at understanding how the performance and the phenology of the invasive legume Lupinus polyphyllus vary with latitude. We sampled data across a >2000 km latitudinal gradient from Central to Northern Europe. We quantified variation in phenology of flowering and fruiting of L. polyphyllus using >1600 digital photos of inflorescences from 220 individual plants observed weekly at 22 sites. The day of the year at which different phenological phases were reached, increased 1.3–1.8 days per degree latitude, whereas the growing degree days (gdd) required for these phenological phases decreased 5–16 gdd per degree latitude. However, this difference disappeared, when the day length of each day included in the calculation of gdd was considered. The day of the year of the earliest and the latest climatic zone to reach any of the three studied phenological phases differed by 23–30 days and temperature requirements to reach these stages differed between 62 and 236 gdd. Probably, the invasion of this species will further increase in the northern part of Europe over the next decades due to climate warming. For invasive species control, our results suggest that in countries with a large latitudinal extent, the mowing date should shift by ca. one week per 500 km at sites with similar elevations. [ABSTRACT FROM AUTHOR]

Lupinus polyphyllus is rich in color, making it a well-known horticultural ornamental plant. However, little is known about the genes related to anthocyanin and carotenoid biosynthesis in L. polyphyllus. In this study, transcriptome sequencing was performed on eight different colors of L. polyphyllus. A total of 1.13 billion clean reads were obtained and assembled into 89,124 unigenes, which were then aligned with six databases, resulting in the identification of 54,823 annotated unigenes. Among these unigenes, 76 and 101 were involved in the biosynthetic pathway of carotenoids and anthocyanins, respectively. In addition, 505 transcription factors were revealed, which belonged to the MYB, R2R3-MYB, NAC, bHLH, and WD40 families. A total of 6,700 differentially expressed genes (DEGs) were obtained by comparative transcriptome analysis. Among them, 17 candidate unigenes (four carotenoid genes, seven anthocyanin genes, and six TFs) were specifically up-regulated for one or more colors of L. polyphyllus. Eight representative candidate unigenes were analyzed by qRT-PCR. The findings enrich the transcriptome database of lupine, and provide a rich molecular resource for research on the coloration mechanism of L. polyphyllus. [ABSTRACT FROM AUTHOR]

Intraspecific variation in growth and defence among plant populations can be driven by differences in (a)biotic conditions, such as herbivory and resources. Introduction of species to novel environments affects simultaneously herbivory encountered by a plant and resource availability both directly and via altered competitive environment. Here, we address the question of how growth (leaf mass per area (LMA), plant size) and resistance traits (leaf alkaloids, leaf trichomes, resistance to a generalist snail) vary and covary between native and introduced populations of the garden lupine, Lupinus polyphyllus. We focused specifically on evolved differences among populations by measuring traits from plants grown from seed in a common environment. Plants from the introduced populations were more resistant against the generalist snail, Arianta arbustorum, and they had more leaf trichomes and higher LMA than plants from the native populations. The composition of alkaloids differed between native and introduced populations, with the native populations having more diversity in alkaloids among them. Resistance was positively associated with plant size and LMA across all populations. Other trait associations differed between native and introduced areas, implying that certain trade-offs may be fundamentally different between native and introduced populations. Our results suggest that, for the introduced populations, the loss of native herbivores and the alterations in resource availability have led to a lower diversity in leaf alkaloids among populations and may facilitate the evolution of novel trait optima without compensatory trade-offs. Such phytochemical similarity among introduced populations provides novel insights into mechanisms promoting successful plant invasions. [ABSTRACT FROM AUTHOR]

A weed plant of the species Lupinus polyphyllus was found in settlements located on the territory of the natural park “Vepssky forest” of the Leningrad region. The plant is known as a transforming species in the southern regions of Russia. The distribution of L. polyphyllus within the Vepsian forest occurs not due to the migration of the species to the north along transport routes, as one might assume, but due to flight from gardens and the formation of spontaneous invasive populations. The goal of the study was to identify the molecular and biochemical characteristics of garden lupine at the northern borders of its secondary range. To interpret the obtained data, the task of the study also included a comparison of intraspecific variability between the “old” invasive populations (in Finland and Central Russia, studied by us earlier) and the “new” naturalizing population of L. polyphyllus in the Vepsian forest. The search for L. polyphyllus localities in the territory of the Vepssky Les natural park was carried out by the route method with geobotanical descriptions of experimental sites (5 m × 5 m). DNA was isolated from eight herbarium specimens of L. polyphyllus (MHA, LE) and fifteen specimens collected in the territory of the natural park “Vepssky Les”. To reveal the internal structure and phylogenetic relationships in lupine populations, networks of nuclear and chloroplast haplotypes and cluster analysis (UPGMA) with the SplitsTree program were used. The total content of polyphenols and flavonoids in the leaves was determined spectrophotometrically. The low inter-locality variability of ITS indicates that the populations of L. polyphyllus in Central Russia and in the North of Russia (St. Petersburg and Vepsskaya Pushcha) have the same origin. Analysis of the chloroplast intergenic spacer (rpl32–trnL) indicates intrapopulation diversity and suggests the presence of microevolutionary processes near the northern limits of the secondary distribution range of L. polyphyllus. The high content of polyphenols and flavonoids in the leaves reveals the adaptive capabilities of lupine in the studied area. Evidence suggests that a neglected species may soon become invasive, as has already happened in other regions.

Abstract Invaders exert new selection pressures on the resident species, for example, through competition for resources or by using novel weapons. It has been shown that novel weapons aid invasion but it is unclear whether native species co‐occurring with invaders have adapted to tolerate these novel weapons. Those resident species which are able to adapt to new selective agents can co‐occur with an invader while others face a risk of local extinction. We ran a factorial common garden experiment to study whether a native plant species, Anthriscus sylvestris, has been able to evolve a greater tolerance to the allelochemicals exerted by the invader, Lupinus polyphyllus. Lupinus polyphyllus produces allelochemicals which potentially act as a novel, strong selective agent on A. sylvestris. We grew A. sylvestris seedlings collected from uninvaded (naïve) and invaded (experienced) sites growing alone and in competition with L. polyphyllus in pots filled with soil with and without activated carbon. Because activated carbon absorbs allelochemicals, its addition should improve especially naïve A. sylvestris performance in the presence of the invader. To distinguish the allelochemicals absorption and fertilizing effects of activated carbon, we grew plants also in a mixture of soil and fertilizer. A common garden experiment indicated that the performances of naïve and experienced A. sylvestris seedlings did not differ when grown with L. polyphyllus. The addition of activated carbon, which reduces interference by allelochemicals, did not induce differences in their performances although it had a positive effect on the aboveground biomass of A. sylvestris. Together, these results suggest that naïve and experienced A. sylvestris plants tolerated equally the invader L. polyphyllus and thus the tolerance has not occurred over the course of invasion.

Plant invasions are among the key drivers of global biodiversity and ecosystem change. They often cause reductions in native species richness and overall biodiversity. Nitrogen-fixing plants are problematic as they affect soil nutrient availability and outcompete species of nutrient-poor sites. Here we assessed the impacts of the legume Lupinus polyphyllus on species and functional diversity of mountain meadow communities in the UNESCO Biosphere Reserve Rhön. We compared species diversity (richness, evenness and effective species number), functional diversity (functional richness, evenness, divergence and dispersion) and similarity of plots in three characteristic vegetation types (Nardus grassland, mesic and wet mountain hay meadows) between different lupine cover classes. We calculated community weighted means (CWMs) of single plant traits and plotted them against lupine cover classes. The invasion of L. polyphyllus homogenizes vegetation composition since the similarity among plots of the different vegetation types increased with increasing lupine cover. It significantly affected species diversity in terms of richness and effective species number and the functional divergence of the vegetation. The trait set of species occurring together with lupine was shifted towards more competitive trait values. We demonstrate strongly negative impacts of L. polyphyllus on different mountain meadow vegetation types since L. polyphyllus, fosters the growth of competitive species and leads to overall more productive plant communities. [ABSTRACT FROM AUTHOR]

Landscape composition and structure may strongly affect the spread of invasive species in landscapes. Landscape analysis provides a powerful toolset for assessing invasive species invasions over time and for planning control measures. We applied a combination of aerial mapping and landscape analysis to assess the invasion of the legume, Lupinus polyphyllus, in the Rhön UNESCO Biosphere Reserve. The Biosphere Reserve contains different types of large and well-connected grasslands threatened by lupine invasion. We assessed the changes in lupine distribution between 1998 and 2016 in a strictly protected part of the Biosphere Reserve by means of landscape structure analysis. The area invaded by L. polyphyllus doubled from 1998 to 2016. While the number of lupine stands decreased by 25%, stand size on average increased by 300%; stands also became less compact during that period. Furthermore, the degree of invasion of different grassland types changed. In 1998, all investigated grassland types were invaded to equal extents, whereas in 2016, large and well-connected mesic grasslands located close to roads were more heavily invaded than small and remote wet grasslands. Our results show that landscape composition plays an important role for the spread of lupine. Specifically, invasive stand characteristics, such as stand size, form, and connectivity, are crucial for driving the invasion of lupine. Therefore, in addition to landscape composition, invasive stand characteristics should be included in the planning of conservation measures. Overall, aerial mapping combined with landscape analysis provides a cost-effective and practical tool for landscape managers to prioritize invasive control measures. [ABSTRACT FROM AUTHOR]

Invaders exert new selection pressures on the resident species, for example, through competition for resources or by using novel weapons. It has been shown that novel weapons aid invasion but it is unclear whether native species co‐occurring with invaders have adapted to tolerate these novel weapons. Those resident species which are able to adapt to new selective agents can co‐occur with an invader while others face a risk of local extinction. We ran a factorial common garden experiment to study whether a native plant species, Anthriscus sylvestris, has been able to evolve a greater tolerance to the allelochemicals exerted by the invader, Lupinus polyphyllus. Lupinus polyphyllus produces allelochemicals which potentially act as a novel, strong selective agent on A. sylvestris. We grew A. sylvestris seedlings collected from uninvaded (naïve) and invaded (experienced) sites growing alone and in competition with L. polyphyllus in pots filled with soil with and without activated carbon. Because activated carbon absorbs allelochemicals, its addition should improve especially naïve A. sylvestris performance in the presence of the invader. To distinguish the allelochemicals absorption and fertilizing effects of activated carbon, we grew plants also in a mixture of soil and fertilizer. A common garden experiment indicated that the performances of naïve and experienced A. sylvestris seedlings did not differ when grown with L. polyphyllus. The addition of activated carbon, which reduces interference by allelochemicals, did not induce differences in their performances although it had a positive effect on the aboveground biomass of A. sylvestris. Together, these results suggest that naïve and experienced A. sylvestris plants tolerated equally the invader L. polyphyllus and thus the tolerance has not occurred over the course of invasion. [ABSTRACT FROM AUTHOR]

Lupines ( Lupinus spp.) are a common legume found on western U.S. rangelands. Lupinus spp. may contain quinolizidine and/or piperidine alkaloids that can be toxic and/or teratogenic to grazing livestock. Lupinus leucophyllus and Lupinus polyphyllus in particular represent important species in the rangelands of southeast Washington and northeast Oregon. The objectives of this study were to identify the alkaloid profiles of these two species, and to explore the phylogenetic relationship among the different populations using amplified fragment length polymorphism (AFLP) markers. Two chemotypes were found among the 14 accessions of Lupinus leucophyllyus. Chemotype A contained anagyrine, a potent teratogen in cattle, and thus would pose a risk to cause crooked calf syndrome in grazing cattle, while Chemotype B did not contain anagyrine and poses no teratogenic risk. No alkaloids were detected in the L. polyphyllus plants collected in this area. Phylogenetic analysis showed that L. leucophyllus accessions with the same chemotype that were geographically proximal were closely grouped in the cladogram; however, accessions that were geographically proximal that represented different chemotypes did not cluster together. Taken together, these results suggest that for the studied species, chemotype is the principle factor in determining relatedness followed by geography. [ABSTRACT FROM AUTHOR]

Background and Aims Assessing the demographic consequences of genetic variation is fundamental to invasion biology. However, genetic and demographic approaches are rarely combined to explore the effects of genetic variation on invasive populations in natural environments. This study combined population genetics, demographic data and a greenhouse experiment to investigate the consequences of genetic variation for the population fitness of the perennial, invasive herb Lupinus polyphyllus. Methods Genetic and demographic data were collected from 37 L. polyphyllus populations representing different latitudes in Finland, and genetic variation was characterized based on 13 microsatellite loci. Associations between genetic variation and population size, population density, latitude and habitat were investigated. Genetic variation was then explored in relation to four fitness components (establishment, survival, growth, fecundity) measured at the population level, and the long-term population growth rate (λ). For a subset of populations genetic variation was also examined in relation to the temporal variability of λ. A further assessment was made of the role of natural selection in the observed variation of certain fitness components among populations under greenhouse conditions. Key Results It was found that genetic variation correlated positively with population size, particularly at higher latitudes, and differed among habitat types. Average seedling establishment per population increased with genetic variation in the field, but not under greenhouse conditions. Quantitative genetic divergence (QST) based on seedling establishment in the greenhouse was smaller than allelic genetic divergence (F'ST), indicating that unifying selection has a prominent role in this fitness component. Genetic variation was not associated with average survival, growth or fecundity measured at the population level, λ or its variability. Conclusions The study suggests that although genetic variation may facilitate plant invasions by increasing seedling establishment, it may not necessarily affect the long-term population growth rate. Therefore, established invasions may be able to grow equally well regardless of their genetic diversity. [ABSTRACT FROM AUTHOR]

The success of plant invasions may be limited by the availability of propagules and/or of suitable microsites, with microsite availability being affected by, for example, disturbance and interspecific competition. A mechanistic understanding of the contributions of propagule pressure and microsite limitation to plant invasions is therefore required to minimise future invasions. Here, we investigated the relative roles of propagule pressure, the availability of microsites, and their interaction on the establishment of an invasive herb, Lupinus polyphyllus , in two geographic regions representing different climate and growth conditions in Finland (a more productive southern region and a harsher central region). We carried out a field experiment in 14 L. polyphyllus populations, in which we manipulated both propagule pressure and disturbance. In a complementary greenhouse experiment, we manipulated propagule pressure and interspecific competition. Seedling establishment of L. polyphyllus was higher in the more productive southern region than in the harsher central region. The number of L. polyphyllus seedlings increased with increasing propagule pressure regardless of disturbance or interspecific competition. However, the number of L. polyphyllus seedlings per sown seed (relative establishment) tended to decrease with increasing propagule pressure, indicating that the positive effect of propagule pressure on early invasion is partially counteracted by density-dependent mortality at high seed densities. Our results highlight the dominant role of propagule pressure over disturbance and interspecific competition in the establishment of L. polyphyllus , suggesting that the early stage of invasion is limited by the availability of propagules rather than the availability of suitable microsites. [ABSTRACT FROM AUTHOR]

Lupinus polyphyllus, known in New Zealand as Russell lupin, is an herbaceous perennial that is invasive in a number of countries outside its natural range in North America. Knowledge of the optimum timeframe and plant development stage for herbicide application is crucial for any effective and cost-efficient weed-control operation. We examined the development and viability ofL. polyphyllusseeds 7 weeks after foliar application of triclopyr ester herbicide at three stages of seed-pod maturity (immature, intermediate and mature). The proportion of seeds in each seed-development class and each seed/seedling viability class from mature pods on treated plants was similar to that in the untreated control. Pods sprayed at immature and intermediate development stages had significantly lower proportions of fully developed seeds (average 82.1% lower) and produced significantly lower proportions of viable seedlings (average 92.7% lower). Our findings indicate thatL. polyphylluscontrol using triclopyr ester can be extended beyond the pre-reproductive stage to include plants bearing up to intermediate-stage seed pods without substantial production of viable seedlings or viable but dormant seed. [ABSTRACT FROM AUTHOR]

Some native species present challenges for efficient propagation and production due to small seed size, low germination and establishment rates, as well as sensitivity to transplanting. In the Pacific Northwest, 3 species, Spiraea douglasii Hook. (Rosaceae), Holodiscus discolor (Pursh) Maxim. (Rosaceae), and Lupinus polyphyllus Lindl. (Fabaceae), are grown as container or bareroot (plug+1) stocktypes at the Coeur d'Alene Forest Service nursery in northern Idaho using Jiffy plugs for ease of germination, growth, and planting. We have found that this method overcomes production challenges for small-seeded bareroot and container plants and shortens time to seed harvest for seed increase needs. [ABSTRACT FROM AUTHOR]

The community-level impacts of invasive plants are likely to vary depending on the character of native species of the target communities and their ability to thrive within the stands of the dominant alien invader. Therefore, I examined the response of native species richness to the cover of the dominant alien Lupinus polyphyllus in two distinct invaded ranges: Czech Republic (Central Europe) and New Zealand. I compared the relation between native species richness and the cover of the dominant alien L. polyphyllus with that in its native range, Pacific Northwest, USA. In the native range, I found no response of native species richness to the cover of L. polyphyllus. In the Czech Republic (central Europe), the richness of native species related to it negativelly, but the relation was only marginally significant. Contrary to that, the richness of species native to New Zealand related to the cover of L. polyphyllus strongly negatively and the negative relation was significantly stronger than that of species native to Europe. Of the two invaded ranges, species native to New Zealand have been documented to be much more vulnerable to the conditions associated with the invasion and dominance of L. polyphyllus, compared to species native to central Europe. This principle has been shown both across these two invaded ranges and in New Zealand, where the aliens of european origin successfully coexist with the dominant invasive alien L. polyphyllus. Similarly, species in the native range of L. polyphyllus showed no relation to its cover, indicating their ability to thrive even in dense stands of this dominant species. [ABSTRACT FROM AUTHOR]

Intraspecific variation in seed size may result from life-history constraints or environmental conditions experienced. This variation in seed size is likely to affect the early stage of invasion as seed size may contribute to the success or failure of population establishment. However, only a few studies have examined seed size variability and its causes and consequences for invaders so far. Using the invasive herb Lupinus polyphyllus, we estimated seed mass variation within and among 39 populations from two different geographic regions in a part of the invaded range. We empirically and experimentally evaluated the effect of seed number and environmental conditions (e.g. geographic region, habitat type, intraspecific competition) on seed mass, emergence and seedling performance. Seed mass varied threefold, being largest among individual plants within populations and smallest among populations. Variation in seed mass was neither related to seed number nor the environmental conditions examined, but led to differences in offspring performance, with emergence and seedling size increasing with seed mass. Larger L. polyphyllus seeds were better establishers than smaller seeds regardless of environmental conditions, indicating that the success of L. polyphyllus invasions is likely to depend positively on seed mass. Our results suggest that some plant species such as the invasive L. polyphyllus may not show an adaptive response in seed mass to resources or environmental conditions, which may partly explain their ability to colonise a range of different habitats. [ABSTRACT FROM AUTHOR]

The aim of this work was to examine H-ATPase hydrolytic activity and the stability of transmembrane electrochemical gradients in membrane vesicles isolated from seedlings and leaf cells of an invasive Washington lupine ( Lupinus polyphyllus Lindl.) and compare them with non-invasive yellow lupine ( Lupinus luteus L.). Temperatures of 25 and 30°C, keeping in mind possible climate warming, were used. For harder stress conditions, short term cold treatment (−8°C in vivo) was used. Plasmalemma-, tonoplast-, and endoplasmic reticulum-enriched membrane fractions were obtained from a sucrose density gradient and identified. Differences in ATPase hydrolytic activity were significant only between lupine species and were more obvious in plasmalemma-enriched fractions. Preincubation of seedlings and leaves at −8°C for 15 min to 24 h showed that microsomic fraction membranes of invasive lupine were more stable (according to Na-diffusive potential) at low temperature compared to yellow lupine ones. The level of transmembrane electrochemical potential, mainly evoked by ATP-dependent active proton transport, was almost equal in both lupine species. Supposedly, the cells of invasive lupine are able to maintain transmembrane electrochemical potential by the employment of lower hydrolytic activity of H-ATPase, thus saving energy for growth. [ABSTRACT FROM AUTHOR]

Lupinus polyphyllus is rich in color, making it a well-known horticultural ornamental plant. However, little is known about the genes related to anthocyanin and carotenoid biosynthesis in L. polyphyllus . In this study, transcriptome sequencing was performed on eight different colors of L. polyphyllus . A total of 1.13 billion clean reads were obtained and assembled into 89,124 unigenes, which were then aligned with six databases, resulting in the identification of 54,823 annotated unigenes. Among these unigenes, 76 and 101 were involved in the biosynthetic pathway of carotenoids and anthocyanins, respectively. In addition, 505 transcription factors were revealed, which belonged to the MYB , R2R3-MYB , NAC , bHLH , and WD40 families. A total of 6,700 differentially expressed genes (DEGs) were obtained by comparative transcriptome analysis. Among them, 17 candidate unigenes (four carotenoid genes, seven anthocyanin genes, and six TFs) were specifically up-regulated for one or more colors of L. polyphyllus . Eight representative candidate unigenes were analyzed by qRT-PCR. The findings enrich the transcriptome database of lupine, and provide a rich molecular resource for research on the coloration mechanism of L. polyphyllus ., Competing Interests: The authors declare there are no competing interests., (©2022 Gao et al.)

The effects of plant invasions on plant communities are often assessed at a few sites or in a particular type of habitat, while studies in different habitat types are scarce. We investigated plant communities in the presence and absence of the invasive ornamental herb, Lupinus polyphyllus, in four habitat types: meadow, forest, road verge and wasteland, in two geographic regions by comparing vascular plant species richness, vegetation structure based on species traits, community composition and the reproductive success of native plant species at invaded and non-invaded locations. The invader was associated with declines in the number of vascular plant species in all habitat types but was unassociated with differences in plant community composition in terms of species identity or species relative cover. However, sites with large lupin invasions (≥1,000 m) contained fewer vascular plant species, a larger proportion of clonal species and more lighter-seeded species than sites with small invasions. The reproductive output of native plants varied across habitats from declines to increases in the presence of L. polyphyllus, and depended on species status (meadow, non-meadow species) rather than species identity, with meadow species generally showing an increase in the reproductive output in the presence of the invader. Overall, our results demonstrate that lupin invasions are associated with declines in local plant species richness across habitats. Although we did not detect systematic differences in species composition between invaded and non-invaded locations, species with particular traits may still be more persistent in invaded plant communities than others. [ABSTRACT FROM AUTHOR]

Morphological characters of floral element of Lupinus polyphyllus at different stages of development are retraced. It is revealed that a blossom dust is emitted twice. The garden form hardly differs from invasive populations, except for a slightly larger size. The size and the form of pollen grains correlate with the color of florets. The pollen fertility of pink florets is dependent on its position in an inflorescence-the main or lateral parts. L. polyphyllus has the competitive superiority over closely related L. angustifolius in a greater number of florets in an inflorescence and specificity of androecium which allows each flower to emit a blossom dust twice. [ABSTRACT FROM AUTHOR]

Abstract: The alkaloid profile of white lupine Lupinus albus L. cv. ‘Boros’ and ‘Butan’, narrow-leaf lupine L. angustifolius L. cv. ‘Bojar’, ‘Graf’, ‘Karo’, ‘Mirela’, and ‘Sonet’, yellow lupine L. luteus L. cv. ‘Dukat’, ‘Parys’, ‘Perkoz’, and ‘Talar’, and wild species big-leaf lupine L. polyphyllus Lindl. was studied. The pea aphid Acyrthosiphon pisum Harris probing behaviour was monitored using the electrical penetration graph (EPG) technique. Four types of feeding behaviour were recognized. First, on highly acceptable L. luteus cv. ‘Dukat’, the total and mean duration of probing, time to reach phloem phase, and the duration of the first phloem phase were comparable to those in aphids on control plant Pisum sativum. Second, on partially acceptable L. luteus cv. ‘Talar’, pathway activities were slightly impeded, and the probes were more numerous and slightly shorter than on control plants. The phloem phase occurred sporadically, and feeding was terminated early after a brief period of ingestion. Third, on unpalatable but acceptable L. angustifolius cv. ‘Bojar’ and ‘Sonet’, and L. luteus cv. ‘Parys’ and ‘Perkoz’, the probes were numerous and usually epidermal. The phloem phase occurred rarely, and when it did, it was short and consisted mainly of watery salivation. Finally, on unacceptable L. albus cv. ‘Boros’ and ‘Butan’, L. angustifolius cv. ‘Graf’, ‘Karo’, ‘Mirela’, and L. polyphyllus, total probing time and individual probes were short and phloem phase did not occur. Eighteen alkaloids were identified: one piperidine alkaloid (ammodendrine), one indole (gramine), ten quinolizidine alkaloids (one tricyclic and nine tetracyclic compounds), and six esters. All lupine varieties that contained lupanine, its derivatives and especially their esters appeared to be unacceptable to the pea aphid, independent of the total concentration of any specific lupanine alkaloid. In contrast, sparteine and its derivatives did not seem to affect aphid probing significantly. [Copyright &y& Elsevier]

Prioritisation of high-impact species is becoming increasingly important for management of introduced species ('neobiota') because of their growing number of which, however, only a small fraction has substantial impacts. Impact scores for prioritising species may be affected by the type of effect model used. Recent studies have shown that environmental co-variation and non-linearity may be significant for effect models of biological invasions. Here, we test for differences in impact scores between simple and complex effect models of three invasive plant species (Heracleum mantegazzianum, Lupinus polyphyllus, Rosa rugosa). We investigated the effects of cover percentages of the invasive plants on species richness of invaded communities using both simple linear effect models ('basic models') and more complex linear or nonlinear models including environmental co-factors ('full models'). Then, we calculated impact scores for each invasive species as the average reduction of species richness predicted by basic and full effect models. All three non-native species had negative effects on species richness, but the full effect models also indicated significant influence of habitat types. Heracleum mantegazzianum had uniform linear effects in all habitats, while effects of L. polyphyllus interacted strongly with habitat type, and R. rugosa showed a marked non-linear relationship. Impact scores were overestimated by basic effect models for H. mantegazzianum and R. rugosa due to disregard of habitat effects and non-linearity, respectively. In contrast, impact of L. polyphyllus was underestimated by the basic model that did not account for the strong interaction of invader cover and habitat type. We conclude that simple linear models will often yield inaccurate impact scores of non-native species. Hence, effect models should consider environmental co-variation and, if necessary, non-linearity of the effects of biological invasions on native ecosystems. [ABSTRACT FROM AUTHOR]

ivated carbon (AC) is widely used in ecological studies to elucidate the role of allelopathic substances in interspecific plant competition. However, by adsorbing chemical signalling compounds AC may also have negative effects on plants with symbiosis partners such as arbuscular mycorrhizal fungi and rhizobia. Here we test whether addition of AC has detrimental effects on the mycorrhizal root colonization of the native forb Plantago lanceolata and the exotic legume Lupinus polyphyllus, the nodulation of L. polyphyllus, and the nutrient uptake and growth of the plants growing in intra- and interspecific competition. Allelopathic effects probably occurred in the germination and seedling establishment phase when P. lanceolata suffered from the presence of L. polyphyllus. However, this negative effect of L. polyphyllus on P. lanceolata was not ameliorated by AC addition. AC negatively affected L. polyphyllus root biomass in week 4, and root and shoot biomass of P. lanceolata in week 9 of the experiment; both effects were independent of the presence and absence of the competing plant species. Mycorrhizal root colonization of both plant species was reduced in the presence of AC, although the effect tended to be stronger for L. polyphyllus. No significant effect of AC on the nodulation of L. polyphyllus was detected. P. lanceolata was the superior competitor and led to reduced biomasses of L. polyphyllus in interspecific competition. We conclude that AC can reduce the mycorrhization and performance of plants which may lead to changes in interspecific competition without the involvement of allelopathy. Contrary to former studies the AC used in our study did not enhance the nutrient availability for the plants, but reduced plant growth and mycorrhization. We suggest that the nutrient properties of the used AC are of crucial importance for the direction and the mechanisms of the effects and should always be reported. [ABSTRACT FROM AUTHOR]

Monosaccharide transporter (MST) genes of Lupinus polyphyllus and L. albus were cloned, expressed and characterised. The isolation and functional characterisation of a cDNA clone and its corresponding genomic clone of a sugar transporter from L. polyphyllus (LpSTP1) is reported. Phylogenetic comparison of the nucleic and amino acid sequences showed the highest similarity to the AtSTP1 gene from Arabidopsis thaliana, which encodes a high affinity sugar transporter. The similar topology as well as the substrate specificity and expression pattern of LpSTP1 encoded protein additionally support the high similarity to the AtSTP1 gene product. The 1,590 bp LpSTP1 cDNA clone was heterologously expressed in yeast resulting in a fully functional specific sugar transporter. This transformation restored the viability of a yeast deletion mutant, which is devoid of all intrinsic MSTs and thus unable to take up and grow on hexose-containing media. The LpSTP1 protein is postulated to be a high-affinity MST since it supported growth best on media containing 0.2% hexose. Tissue-specific expression of LaSTP1 in L. albus was assayed by real-time PCR, which revealed that the lupin STP1 is mainly expressed in flower buds, flowers and young leaves. The results suggest that the main role of LaSTP1 is to catalyse monosaccharide import in sink tissues to meet increased carbohydrate demand during plant development. [ABSTRACT FROM AUTHOR]

1 Assembly rules are broadly defined as any filter imposed on a regional species pool that acts to determine the local community structure and composition. Environmental filtering is thought to result in the formation of groups of species with similar traits that tend to co-occur more often than expected by chance alone, known as Beta guilds. At a smaller scale, within a single Beta guild, species may be partitioned into Alpha guilds – groups of species that have similar resource use and hence should tend not to co-occur at small scales due the principle of limiting similarity. 2 This research investigates the effects of successional age and the presence of an invasive exotic species on Alpha and Beta guild structuring within plant communities along two successional river terrace sequences in the Waimakariri braided river system in New Zealand. 3 Fifteen sites were sampled, six with and nine without the Russel lupin ( Lupinus polyphyllus), an invasive exotic species. At each site, species presence/absence was recorded in 100 circular quadrats (5 cm in diameter) at 30-cm intervals along a 30-m transect. Guild proportionality (Alpha guild structuring) was tested for using two a priori guild classifications each containing three guilds, and cluster analysis was used to test for environmental structuring between sites. 4 Significant assembly rules based on Alpha guild structuring were found, particularly for the monocot and dicot guild. Guild proportionality increased with increasing ecological age, which indicated an increase in the relative importance of competitive structuring at later stages of succession. This provides empirical support for Weiher and Keddy's theoretical model of community assembly. 5 Lupins were associated with altered Alpha and Beta guild structuring at early mid successional sites. Lupin-containing sites had higher silt content than sites without lupins, and this could have altered the strength and scale of competitive structuring within the communities present. 6 This research adds to the increasing evidence for the existence of assembly rules based on limiting similarity within plant communities, and demonstrates the need to incorporate gradients of environmental and competitive adversity when investigating the rules that govern community assembly. [ABSTRACT FROM AUTHOR]

Wild tobacco growing in a field near experimentally clipped sagebrush became more resistant to herbivores than tobacco plants near unclipped sagebrush. Here, we asked whether eavesdropping on sagebrush is a common phenomenon. Do three other forb species that grow near sagebrush experience less herbivory when their sagebrush neighbors are clipped experimentally? Similarly, we asked if sagebrush becomes more resistant when conspecific neighbors are clipped. We found that levels of damage to leaves of lomatium, lupine, and valerian were unaffected by experimental clipping of neighboring sagebrush. These three forbs share many generalist herbivores with sagebrush, as does wild tobacco. Sagebrush individuals near clipped neighbors experienced approximately half the leaf loss to herbivores over the summer as plants with unclipped neighbors. Most of this herbivory was caused by generalist grasshoppers and mammals. Success of a chrysomelid beetle that specializes on sagebrush was unaffected by clipping neighboring sagebrush. These results indicate that communication between sagebrush and tobacco is not a unique phenomenon since sagebrush also responded when neighbors were clipped. In contrast, other plant species generally respond less strongly to cues than does tobacco, indicating that not all species respond to relatively high levels of jasmonate in their environments. Eavesdropping may be restricted to those species, like tobacco, that are strongly inducible and to those species, like sagebrush, that release high concentrations of cues. [ABSTRACT FROM AUTHOR]

Biotin is an essential micronutrient needed but not synthesized by animals. We performed a field experiment where we measured biotin levels and the community composition of nematodes in the rhizosphere of two legumes, the native Lotus corniculatus and an invasive alien, Lupinus polyphyllus. The relative abundance of root-feeding nematodes and the biotin concentration in nematodes were lower in the rhizosphere of the alien. We are the first to show that biotin levels in indigenous nematode populations are associated with the host plant species, and that the difference may affect nematode community. [ABSTRACT FROM AUTHOR]