Effects of climatic factors on Fallopia japonica s.l. seedling establishment: evidence from laboratory experiments

The highly invasive Fallopia complex (Fallopia japonica s.l., Fallopia japonica var. japonica, Fallopia sachalinensis and the hybrid Fallopia × bohemica) is known to spread primarily vegetatively by rhizomes and stem fragments in its adventive range. Sexual reproduction has rarely been shown in Europe, despite a high production of germinable seeds. To elucidate this contradiction the aim of the present study was to determine which environmental conditions prevent seed germination or seedling establishment. Several laboratory experiments were conducted focusing on water and light supply as well as the influence of frost. According to morphological and cytological analyses of the parental plants paired with morphological analyses of the seedlings, the seeds used for these experiments were assumed to belong predominantly to the hybrid Fallopia × bohemica. Laboratory experiments revealed that seedling establishment is hampered by severe drought or strong late frost, which are, however, rare events in the study region and could not be crucial in explaining the rarity of seedlings in the field. We discuss the possible role of weather conditions that might lead to a loss of seeds or seedlings under field conditions and include a comparison of several other cited studies carried out in the adventive range of Fallopia japonica s.l. that observed either the presence or the absence of Fallopia seedlings in the field.     

Winter hosts of Aphalara itadori (Hemiptera: Psyllidae), a classical biological control agent of Fallopia japonica (Polygonaceae), in the UK

This study suggests that lack of suitable overwintering sites will not be a limiting factor to the establishment of Aphalara itadori, a classical biological control agent of Fallopia japonica, since it was able to overwinter on a common conifer species as well as non-conifers hosts.     

Duration of freezing necessary to damage the leaves of Fallopia japonica (Houtt.) Ronse Decraene

Fallopia japonica (Houtt.) Ronse Decraene is an invasive plant species that introduces economic, social, and environmental stresses. After observing frost damage to F. japonica plants in the field, we exposed leaves of F. japonica and a native species (Acer saccharum Marshall) to freezing temperatures in the laboratory and compared their net photosynthetic rate to that of fresh leaves. In both species, the net photosynthetic rate of leaves frozen for 0.5 h or for 1 h were not significantly different from each other but were both significantly less than that of fresh leaves. Fresh leaves of F. japonica had a higher net photosynthetic rate than those of A. saccharum, but the relationship was reversed in all freezing treatments. Frozen leaves of F. japonica contained microscopically visible frost lenses, which revealed the mechanism of the damage. These results quantify how quickly F. japonica is damaged by freezing conditions and suggest that minimum vernal temperatures may limit its range expansion.     

Gallerucida bifasciata (Coleoptera: Chrysomelidae), a potential biological control agent for Japanese knotweed (Fallopia japonica)

Host specificity of foreign natural enemies are becoming more and more critical in classical biological control programs, as concerns about potential risk from introduced biocontrol agents have been increasing recently. Understanding the insect's fundamental and ecological host ranges is the first step in determining the potential for introduction of an insect to control invasive plants. Japanese knotweed, Fallopia japonica (Houttuyn) Ronse Decraene (Polygonaceae) is an invasive weed in the United States and Europe. A leaf beetle, Gallerucida bifasciata (Coleoptera: Chrysomelidae) is an important natural enemy attacking this plant in Asia. However, its host range records were ambiguous. This study examined the beetle's host specificity through a set of choice and no-choice tests in the laboratory and field in its native China. Gallerucida bifasciata larvae were able to complete development on seven of 87 plant species in larval development tests, while adults fed and oviposited on 10 plants in no-choice tests. Multiple choice tests showed adults strongly preferred Fallopia japonica, Persicaria perfoliata (L.) H. Gross and Polygonum multiflorum Thunb over all other plants. Open field tests and field surveys further revealed that these three species were in its ecological host range. The results of this study suggest that G. bifasciata is a potential promising agent for control of Japanese knotweed in the United States and Europe, although additional host specificity tests and risk assessment should be completed.     

Morphological and chromosomal variation in Fallopia section Reynoutria (Polygonaceae) in Korea

We have examined morphological and chromosomal variation inFallopia sect.Reynoutria in Korea to clarify their taxonomic identities and to determine whether their morphological variability is associated with ploidy levels. Principal components analysis (PCA) of individuals from 21 populations, using major distinguishing characters, revealed the presence of four major entiries of sect.Reynoutria in Korea; these includeF. sachalinensis, F. japonica var.japonica, F. forbesii, and the Nonsan population consisting of presumed hybrids. Based on morphology, it is hypothesized that the Nonsan population was probably derived from multiple hybridization events involving the three named taxa. The results also indicate thatF. forbesii is distinct fromF. japonica var.japonica. Polyploidy is more prevalent in sect.Reynoutria than has been previously recognized.Fallopia sachalinensis in Korea occurs as dodecaploids with 2n=132; our count is the first dodecaploid count for the species, and represents the highest chromosome number known in the genus.Fallopia japonica var.japonica occurs as tetraploids (2n=44), hexaploids (2n=66), and octoploids (2n=88), whileF. forbesii occurs as hexaploids (2n=66) and octoploids (2n=88); our counts appear to be the first reported chromosome numbers forF. forbesii. Morphological analysis indicates that there is no apparent correlation between the ploidy levels in these taxa and the morphological characters that we have considered in this study except that the tetraploids ofF. japonica var.japonica tend to have somewhat thicker leaves.     

From plant traits to invasion success: Impacts of the alien Fallopia japonica (Houtt.) Ronse Decraene on two native grassland species

Alien invasive plants threaten biodiversity, productivity and ecosystem functioning throughout the world. We examined the effect of Fallopia japonica on two native grassland species (Trifolium repens, Lolium perenne). We hypothesized that its negative effects on the native species are dependent on three mechanisms: (i) allelochemicals released and accumulated in soil with a history of invasion, (ii) altered soil biota and (iii) direct resource competition. We measured the response of the native species as the difference in their functional traits when grown under the three conditions. Our results demonstrate that neither allelochemicals nor soil biota from soil with history of F. japonica invasion had measurable effects on either species. Competition with the invader strongly reduced height, biomass and specific leaf area (SLA) of T. repens, while it had a lower effect on L. perenne. Furthermore, our results reveal that F. japonica took advantage of a positive plant–soil and plant–plant interaction. The results show that the prominent mechanism underpinning the invasion success of F. japonica in the grassland was the direct resource competition. This prominent role is also confirmed by the significant interactions between competition, allelochemicals and soil biota from soils with history of invasion of F. japonica on SLA of the native species.     

Seed characteristics and germination limitations in the highly invasive <Emphasis Type="Italic">Fallopia japonica</Emphasis> s.l. (Polygonaceae)

The species in the Japanese knotweed complex (Fallopia japonica s.l. and its hybrids) are among the most invasive plants on earth. Their expansion and reproduction in the introduced range have been mostly due to vegetative reproduction, but observations of low seedling numbers and hybridization processes exist. Knowledge of factors affecting germination characteristics is essential if the risk of sexual reproduction is to be assessed, and its impact on the ability of the species to spread and adapt to different environments. This study aims to examine the germination success of Fallopia japonica s.l. seeds of different ages, quality, and storage conditions. Irrespective of age and even after natural overwintering in the soil, seeds germinated quite well (48–79%). Ungerminated seeds collected in autumn of 2008 were characterized by a low weight/length ratio (W/L), low nutrient concentrations, and a greenish tepal coloration. These differences may be due to alternative male taxa participating in the pollination process. Spring collected seeds were subject to strong predation by birds. In contrast to the high germination observed under laboratory conditions, seed germination or early establishment in the field was inhibited and only a few seedlings were observed. Although the factors that inhibit the establishment of mature seeds in the field remain unknown, there is a clear risk that sexual reproduction could gain importance in the future as a result of changing environmental conditions or genetic adaptation. Not only would this facilitate expansive dispersal by wind, but it might also increase the potential for further adaptation of the species complex.     

The potential role of polyploidy and hybridisation in the further evolution of the highly invasive <Emphasis Type="Italic">Fallopia</Emphasis> taxa in Europe

Japanese knotweed s.l. comprises Fallopia japonica, F. sachalinensis, F. × bohemica and any F2s or backcrosses. The parental taxa were introduced from the East to the West as garden ornamentals in the nineteenth century, and soon spread beyond the confines of the garden to become widespread and persistent weeds. Since only female F. japonica var. japonica was introduced, its impressive spread has occurred solely by vegetative means. However, the initial lack of genetic variability has been complemented by an extensive series of hybridisations in the adventive range. We examine the history, spread, reproductive biology and ecological impact of these species in the West. The role and importance of polyploidy and hybridisation in their invasion of the West is discussed, as are the implications of these factors for the potential further evolution of the group.     

Haplotypes of <Emphasis Type="Italic">Fallopia</Emphasis> introduced into the US

In the US, clonal growth of Fallopia japonica, Fallopia sachalinensis and their hybrid Fallopia x bohemica (Polygonaceae) is prominent, yet sexual reproduction and hybridization contribute to the genetic complexity of swarms. The contribution to this diversity from multiple introductions is unknown. Using 800 bp of the non-coding chloroplast marker accD–rbcL, we compared 21 Japanese haplotypes with 46 US samples from 11 states, 2 Canadian samples, and 6 European samples from 4 countries, in order to investigate if there were repeated introductions from Asia. While most North American and all European haplotypes accessions in our collection matched a single widespread haplotype, we identified 8 other haplotypes. Three haplotypes of F. japonica (including the widespread haplotype) and one F. sachalinensis matched previously identified Japanese haplotypes, supporting the hypothesis of multiple introductions in the US. Five additional US haplotypes were detected once. Four of these differed from Japanese haplotypes by one single nucleotide polymorphism (SNP), possibly indicating a recent in situ change. The fifth haplotype represents a garden cultivar, which differed from all F. japonica haplotypes. It therefore appears that the US genetic diversity of these taxa has three sources: intra-specific reproduction, inter-specific reproduction, and multiple sources of introduction.     

Identification of quantitative trait loci associated with germination using chromosome segment substitution lines of rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.)

Rapid and uniform seed germination under diverse environmental conditions is a desirable characteristic for most crop plants, such as rice, wheat, and maize. However, the genetic base of the variations in the rate of germination is not well understood. In this study, quantitative trait loci (QTL) for germination rate were mapped with a set of 143 chromosome segment substitution lines (CSSL) each contains a small genomic fragment from a japonica variety Nipponbare in the uniform genetic background of an indica variety Zhenshan97. Nine CSSL showed significantly lower germination rate than that in Zhenshan97. Four germination-related QTL were identified located on chromosomes 2, 5, 6 and 10, at which all japonica alleles decreased germination rate. By using the CSSL-derived F2 population, a major QTL (qGR2) on chromosome 2 was confirmed, and delimited to a 10.4 kb interval containing three putative candidate genes, of which OsMADS29 was only expressed preferentially in the seed. These results would facilitate cloning of the major gene that affects germination rate, and provide an insight into the genetic basis of germination.     

Host specificity of <Emphasis Type="Italic">Euops chinesis</Emphasis>, a potential biological control agent of <Emphasis Type="Italic">Fallopia japonica</Emphasis>, an invasive plant in Europe and North America

Fallopia japonica (Houttuyn) Ronse Decraene (Polygonaceae) is a serious invasive weed in North American and Europe. In its native China, a leaf-rolling weevil, Euops chinesis (Coleoptera: Attelabidae) was found attacking F. japonica in the field. No-choice tests, multiple-choice tests, open field tests and field surveys were conducted as a measure of its host specificity. Forty-six plant species were selected from 17 families for host range testing, among which, six species, F. multiflora, F. japonica, Persicaria perfoliata, Rumex acetosa, R. japonicus and R. aquaticus, were exposed to adults in no-choice tests. However, larvae could only develop successfully on F. japonica, and this plant appeared to be the only host in the field, suggesting the weevil is host-specific. As larval development appears to depend on a fungus in the leaf rolls, the insect–fungus mutualism and risks including host specificity of the fungus should be evaluated before the insect’s introduction.     

Climate and the distribution of Fallopia japonica: use of an introduced species to test the predictive capacity of response surfaces

Abstract. The relationship between present climate and the distribution in Europe of the aggressively invasive exotic Fallopia japonica is described by fitting a response surface based on three bioclimatic variables: mean temperature of the coldest month, the annual temperature sum > 5 °C, and the ratio of actual to potential evapotranspiration. The close fit between the observed and simulated distributions suggests that the species' European distribution is climatically determined. The response surface also provides a simulation of the extent of the area of native distribution of F. japonica in Southeast Asia that is generally accurate, confirming the robustness of the static correlative model upon which it is based. Simulations of the potential distribution of F. japonica under two alternative 2 x CO₂ climate change scenarios indicate the likelihood of considerable spread into higher latitudes and possible eventual exclusion of the species from central Europe. However, despite the robustness of the response surface with present-day climate, the reliability of these simulations as forecasts is likely to be limited because no account is taken of the direct effects of CO₂ and their interaction with the species' physiological responses to climate. Similarly, no account is taken of the potential impact of interactions with ‘new’ species as ecosystems change in composition in response to climate change. Nevertheless, the simulations indicate both the possible magnitude of the impacts of forecast climate changes and the regions that may be susceptible to invasion by F. japonica.     

Impacts of the invasive plant <Emphasis Type="Italic">Fallopia japonica</Emphasis> (Houtt.) on plant communities and ecosystem processes

Fallopia japonica (Japanese knotweed) invades riparian areas and roadsides in New England. This large clonal species drastically alters the appearance of habitats by forming highly productive near-monocultures. To understand how these invasions affect ecosystem processes in New England, we quantified the impacts of F. japonica on species diversity, primary productivity, and nitrogen cycling at five locations in central Massachusetts, USA. In stands of F. japonica and in adjacent uninvaded areas, we recorded the cover of each plant species and measured the aboveground biomass and nitrogen (N) concentrations in plants, along with N retranslocation from F. japonica leaves and several soil characteristics. In addition, we severed rhizomes of peripheral F. japonica shoots to determine if clonal integration contributes to the species’ rapid spread and dominance. Stands of F. japonica had lower species diversity, but greater aboveground biomass and standing N than uninvaded areas. Nitrogen and carbon concentrations in biomass and N mineralization rates in soil did not differ between stands and adjacent areas. Rhizome severing temporarily reduced growth of F. japonica, suggesting that retranslocation of photoassimilates and/or nutrients between shoots via rhizomatal connections may maximize stand level growth rates and facilitate dominance by F. japonica.     

Seasonal patterns of partitioning and remobilization of 14C in the invasive rhizomatous perennial Japanese knotweed (Fallopia japonica (Houtt.) Ronse Decraene)

Resource partitioning between shoot growth, storage and reproduction is poorly understood in many clonal plant species. This study documents seasonal patterns of growth, ¹⁴C-labelled photoassimilate distribution and remobilization in the invasive rhizomatous species Fallopia japonica (Japanese knotweed). Biomass accumulation above- and below-ground in F. japonica was rapid. By September, rhizome biomass had increased 18-fold from the initial harvest in May (representing 48% of total plant biomass) and this was maintained over winter. Patterns of ¹⁴C allocation from F. japonica shoots labelled at different times of year show that as the season progressed, the rhizomes became an increasingly important sink for current assimilate (the percentage of ¹⁴C recovered from rhizomes was 35% in August and 67% in September) and the corresponding retention of assimilate by established shoots declined. The percentage of ¹⁴C exported to roots was greatest in August. Relatively little photoassimilate was exported to other shoots on the plant, or to flowers. Recycling of photoassimilate was fairly tight in this species and ¹⁴C fixed by shoots in early May 1999 or September 1999 was remobilized to the rhizome prior to shoot senescence and death. Some of this ¹⁴C was then remobilized to new shoots early the following spring. These characteristics may contribute to the success of F. japonica in colonizing a variety of contrasting habitats, often with serious management implications.     

Establishment of the biological control agent Aphalara itadori is limited by native predators and foliage age

The knotweed psyllid, Aphalara itadori, is a biological control agent for invasive knotweed species in North America and Europe. Initial releases were conducted in Canada in 2014 but establishment has been slow, seemingly as a result of low nymphal survival. We conducted two field experiments in Ontario, Canada, to explore the effects of native predators and the age of knotweed (Fallopia japonica) foliage on nymphal survival in A. itadori. Survival of A. itadori nymphs was significantly reduced on potted plants that were exposed to native predators in the field, compared to plants from which predators were excluded. The number of surviving nymphs was also significantly reduced on older F. japonica foliage, compared to recent regrowth after a summer cutting treatment. We discuss our findings in the context of biological invasion theory and emphasize the potential for increased overlap between the fields of invasion ecology and biological control. Finally, we advocate the use of A. itadori in combination with other control measures as part of an integrated pest management programme, rather than as a solitary measure. Specifically, we recommend that future releases of A. itadori be concentrated shortly after cutting or herbicide treatments in order to maximize the availability of young tender foliage.     

Asexual spread versus sexual reproduction and evolution in Japanese Knotweed s.l. sets the stage for the “Battle of the Clones”

The invasion of NW Europe by a single clone of male-sterile Fallopia japonica var. japonica by purely vegetative reproduction has demonstrated that reproduction by seed and the genetic diversity associated with it may not be an essential feature of plant invasions, at least in the short term. What is less well known is that a significant proportion of the Japanese Knotweed s.l. involved is not F. japonica var. japonica, but the hybrid between it and F. sachalinensis–F. × bohemica. This hybrid is able to backcross to either parent with the potential to replace the missing male F. japonica; by the same process, the hybrid is generating the genetic diversity so conspicuously lacking in F. japonica. In terms of understanding the population structure in a particular country, it is important to be able to identify hybrids and putative back-crosses. We bring together a mixture of published and unpublished information to provide a comprehensive section on morphological and anatomical aids to identification. Regeneration of plants from stem and rhizome fragments varies by taxon, as do responses to control techniques, underlining the importance of correct identification of these plants. In this paper we look closely at this group of taxa, with special emphasis on the role played by hybridization, with an examination of the genetic make up of seedlings produced in the wild or by artificial hybridizations, and the implications that this will have on the future directions of the invasion process.     

Presence of Cry1Ab in the Bt maize – aphid (Rhopalosiphum maidis) – ladybeetle (Propylea japonica) system has no adverse effects on insect biological parameters

Transgenic Bacillus thuringiensis Berliner (Bt) crops receive particular attention because they carry genes encoding insecticidal proteins that might negatively affect non‐target arthropods. Here, laboratory experiments were conducted to evaluate the impact of Cry1Ab‐expressing transgenic maize [5422Bt1 (event Bt11) and 5422CBCL (MON810)] on the biological parameters of two non‐target arthropods, the aphid Rhopalosiphum maidis (Fitch) (Hemiptera: Aphididae) and its predator the ladybeetle Propylea japonica (Thunberg) (Coleoptera: Coccinellidae). In a long‐term assay (three generations), no significant differences were found between R. maidis fed Bt maize and those fed a near‐isogenic line (5422) when individual parameters were compared, including nymph development time, adult longevity, aphid spawning period, and fecundity. No negative effects were detected throughout the life cycle of P. japonica in aphids’ feeding amount, development (nymphs, pupae, adults, and progeny eggs), fecundity, or egg hatching when they preyed on Bt maize‐fed aphids compared with non‐Bt maize treatments. A tritrophic assay revealed that Cry1Ab was highly diluted through the food chain (Bt maize leaves, R. maidis, and P. japonica), as detected by an enzyme‐linked immunosorbent assay (ELISA). In conclusion, although Cry1Ab concentrations in maize leaves increased as the plants developed, Cry1Ab levels were significantly reduced in the aphid R. maidis, and no traces of Cry1Ab were detected in P. japonica preying on Bt maize‐fed aphids. The two hybrids of Bt maize expressing Cry1Ab had no negative effects on the measured biological parameters of the aphid R. maidis or its predator, the ladybeetle P. japonica.     

Single-seeded InDel fingerprints in rice:An effective tool for indica-japonica rice classification and evolutionary studies

Asian cultivated rice(Oryza sativa L.),an important cereal crop worldwide,was domesticated from its wild ancestor 8000 years ago.During its long-term cultivation and evolution under diverse agroecological conditions, Asian cultivated rice has differentiated into indica and japonica subspecies.An effective method is required to identify rice germplasm for its indica and japonica features,which is essential in rice genetic improvements.We developed a protocol that combined DNA extraction from a single rice seed and the insertion/deletion(InDel) molecular fingerprint to determine the indica and japonica features of rice germplasm.We analyzed a set of rice germplasm,including 166 Asian rice varieties,two African rice varieties,30 accessions of wild rice species,and 42 weedy rice accessions,using the single-seeded InDel fingerprints(SSIF).The results show that the SSIF method can efficiently determine the indica and japonica features of the rice germplasm.Further analyses revealed significant indica and japonica differentiation in most Asian rice varieties and weedy rice accessions.In contrast,African rice varieties and nearly all the wild rice accessions did not exhibit such differentiation.The pattern of cultivated and wild rice samples illustrated by the SSIF supports our previous hypothesis that indica and japonica differentiation occurred after rice domestication under different agroecological conditions.In addition,the divergent pattern of rice cultivars and weedy rice accessions suggests the possibility of an endoferal origin(from crop)of the weedy rice included in the present study.     

Systematics of <Emphasis Type="Italic">Mycosphaerella</Emphasis> species associated with the invasive weed <Emphasis Type="Italic">Fallopia japonica</Emphasis>, including the potential biological control agent <Emphasis Type="Italic">M. polygoni-cuspidati</Emphasis>

Native to Japan, Fallopia japonica, most frequently referred to as Japanese knotweed, is a highly problematic invasive weed, particularly in the UK and North America. During surveys for natural enemies of this plant in Japan, two species of Mycosphaerella were collected. One of these was identified as M. polygoni-cuspidati, and is redescribed and neotypified. Causing a damaging leaf spot disease of F. japonica throughout its natural range in Japan, it is absent from the host’s exotic range. The restriction of M. polygoni-cuspidati to F. japonica in its center of origin, together with its severe impact on host fitness, indicates that this is a coevolved natural enemy with high potential as a classical biological control agent for the long-term management of this ecologically and economically important weed. In the field, the fungus has a reduced life cycle, with only spermogonia and pseudothecia (ascomata) being formed. Ascospores are the primary source of infection, and studies show that the mycelium from in vitro cultures is also infective and hyphae penetrate mainly via the stomata. A further, undescribed species of Mycosphaerella co-occurs with M. polygoni-cuspidati, here proposed as the new species M. shimabarensis. Both species have been studied using cultural, morphological and molecular phylogenetic methods.     

Cup plant,an alternative to conventional silage from a LCA perspective

Purpose

The growing awareness of the importance of biodiversity in agroecosystems in increasing and ensuring the supply of biomass has led to heightened interest from governments and farmers in alternative crops. This article assesses one such alternative crop, cup plant (Silphium perfoliatum L.), in terms of the environmental aspects of cultivation for forage production. Many studies have previously focused on cup plant, but so far, this plant has not been assessed using the life cycle assessment (LCA) method.

Materials and methods

This study compares the environmental load of cup plant with the most commonly grown silage crops in Central European conditions—maize—and with another common forage crop—lucerne using LCA. The system boundaries include all the processes from cradle to farm gate and both mass-based (1 ton of dry matter) and area-based (1 ha of monoculture) functional units were chosen for the purposes of this study. The results cover the impact categories related to the agricultural LCAs, and the ReCiPe Midpoint (H) characterization model was used for the data expression, by using SimaPro 9.0.0.40 software.

Results

This study compares the cultivation of cup plant with the most commonly grown silage crop in Central European conditions—maize—and with another common forage crop—lucerne. The paper shows the potential of cup plant to replace conventional silage (maize and lucerne silage mix) with certain environmental savings in selected impact categories, and importantly, while still maintaining the same performance levels in dairy farming as with conventional silage, as already reported in previous publications. For the Czech Republic alone, this would, in practice, mean replacing up to 50,000 ha of silage maize and reducing the environmental load by about tens of percent or more within the various impact categories and years of cultivation.

Conclusion

Cup plant can replace the yield and quality of silage maize, represents a lower environmental load per unit of production and unit of area and generally carries many other benefits. Thus, cup plant is a recommendable option for dairy farming. Given the recent experience and knowledge of the issue, the cup plant can be considered an effective alternative to conventional silage.