Yield of willow (Salix spp.) grown in short rotation coppice mixtures in a long‐term trial

Salix spp. genotypes were grown in random intimate mixtures comprising 5, 10, 15 and 20 components at 3 planting densities (10 000, 15 000 and 20 000 cuttings ha^?1). Planted in 1994/95, plots were harvested every 3 years in 1998/99 (previously reported), 2001/02, 2004/05 and 2007/08. Each individual stool in both mono‐ and mixture plots was weighed. The total yield from mixture plots was consistently higher than the mean of the components in mono‐plots and with only limited exceptions higher than the yield of any of the individual components grown in mono‐plots. This occurred despite the loss of a number of disease‐susceptible genotypes. When a stool died the remaining plants were able to colonise the vacant space and compensate for the loss. There was no clear benefit in increasing the number of components from 10 to 15 or 20 although host diversity is considered to be an important contributor to the effectiveness of a mixture both in disease reduction and yield enhancement. It is argued that the use of Salix spp. mixtures is highly advantageous, even in the absence of any disease pressure, and that mixtures increase the sustainability of a willow plantation.     

Phenological stages of willow (Salix)

The practice of uniform recording of biological plant growth stages or events has long been practiced in agricultural production. In this study the BBCH (Biologishe Bundesanstalt, Bundessortenamt and CHemical Industry) code has been applied to four precocious species of willows to define growth stages important to this group. The studied taxa represent varieties of potential importance in the Floral Industry. A new BBCH code is proposed where the annual cycle of willows is divided into clearly recognisable and easily distinguishable developmental phases which include eight principal stages, 30 secondary stages and six mesostages. Photographs illustrate the physical appearance of select stages. This proposed BBCH code shows a unified approach which may be applied to a large number of Salix species.     

Nitrogen-induced reduction in leaf phenolic level is not accompanied by increased rust frequency in a compatible willow (Salix myrsinifolia)‐Melampsora rust interaction

Phenolic compounds are potential antimicrobial agents in willows ( Salix spp.). However, little is known about their role in willow‐pathogen interactions. Earlier studies have shown that phenolic levels may vary depending on genotype and environmental factors (such as nutrient availability). In the present study, the combined effects of nitrogen fertilization and infection by pathogenic rust ( Melampsora sp.) on phenolics and growth in willow ( salix myrsinifolia ) clones were investigated. Cutting-propagated plants of eight willow clones were subjected to low and optimum levels of nitrogen fertilization and inoculated with rust urediniospores or mock-inoculated with water. The plants were harvested 20 days after the inoculations and the dry weight of the leaves, stems and roots determined. Phenolics were analysed from leaf material using HPLC. The willow clones differed significantly in their phenolic content, rust frequency and phytomass production. Plants under optimum nitrogen availability showed increased shoot phytomass production and reduced root production. Rust infection increased the root phytomass of willow under both fertilization treatments, but had no consistent effect on the shoot growth of the clones. Low nitrogen availability increased the phenolic levels in willow leaves, but the rust frequency did not differ significantly between plants grown at low and optimum levels of nitrogen fertilization. In several clones, rust infection increased the concentrations of individual phenolic compounds, but there were no significant correlations between phenolic concentrations and the level of rust infection. The results indicate that resistance in compatible willow‐rust interactions may not be directly governed by phenolic compounds.     

Host–pathogen relationship between Salix and Melampsora sheds light on the parentage of some biomass willows

The association between willow ( Salix ) and rust ( Melampsora ) is highly specific. Willows named Salix burjatica , S. dasyclados ( S. × dasyclados ) and S. × calodendron are important in renewable-energy plantations in the UK and western Europe. There has been much controversy over their origin, species status and nomenclature. It has been suggested that they have originated from hybridization between. S. caprea , S. viminalis and S. cinerea . In the present work, 59 willow clones were investigated through morphological examination and detached leaf inoculation using willow differentials, for their association, in southwest England, with M. capraearum and three pathotypes of Melampsora epitea (Me-A, B and C). M. capraearum was found on all clones of S. caprea and its hybrids with S. aurita ; Me-A on all S. viminalis clones; Me-B on wild S. cinerea , S. × calodendron , S. × dasyclados 'De Biardii 445' and S . 'Spaethii'; Me-C on all S. burjatica clones and most S. × dasyclados clones. Both M. caprearum and Me-A infected all S. × sericans ( S. caprea × viminalis ) clones and S. × dasyclados 'LA041/03'. We suggest that S. × dasyclados 'LA041/03' should be treated as S. × sericans ( S. caprea × S. viminalis ); S. burjatica , S. dasyclados and S. × dasyclados as synonyms; S. × dasyclados 'De Biardii 445' as S. × calodendron 'De Biardii 445'; and S . 'Spaethii' as S. × calodendron 'Spaethii'.     

Genetic structure and population dynamics of a heteroecious plant pathogen Melampsora larici-epitea in short-rotation coppice willow plantations

Complex life strategies are common among plant pathogens belonging to rust fungi ( Uredinales ). The heteroecious willow rust Melampsora larici-epitea produces five spore stages and alternates on larch ( Larix ). To shed light on the epidemiology of this pathogen, amplified fragment length polymorphisms (AFLPs) were used to determine the genetic diversity and genetic structure of rust samples collected from coppice willow ( Salix ) plantations at three UK sites (LA, CA and MC) over three sampling dates (September 2000, July 2001 and September 2001). Of the total of 819 isolates, 465 were unique AFLP phenotypes and there was a shift in genotype diversity between the two seasons (0.67 in 2000 and 0.87–0.89 in 2001). No phenotypes were common between the two seasons within a site, suggesting that the rust did not overwinter as an asexual stage within plantations. A temporal analysis detected large amounts of genetic drift ( F _S = 0.15–0.26) between the two seasons and very small effective population sizes ( N _e  =  2–3) within sites. These results all point to a new colonization of the plantations by the rust in the second season (2001). The F _ST-analogue values were Φ_CT = 0.121, Weir and Cockerham's θ = 0.086 and the Bayesian estimate θ^B = 0.087–0.096. The results suggest that the sources of inoculum were somewhat localized and the same sources were mainly responsible for disease epidemics in LA and CA over the two seasons. The relatively low F _ST-values among sites (0.055–0.13) suggest the existence of significant gene flow among the three sampled sites.     

Heat shock proteins in willow (Salix viminalis)

The heat shock response in three vegetatively propagated clones of Salix viminalis L. was studied. In the clone 78198, synthesis of a total of 58 proteins was induced or increased by heat shock. Of these proteins, 39 were found in both leaves and callus, 8 only in leaves, and 11 only in callus. The number of heat shock proteins differed between the three clones studied. The molecular weights of the heat shock proteins ranged from 18000 to over 94000. The optimal synthesis of heat shock proteins took place at 37–40°C, but several proteins could be induced at 25–30°C. The synthesis of the majority of the proteins present at a normal growth temperature (20°C) was not completely blocked by the heat shock. More than 12 h was needed for the reappearance of the normal protein synthesis pattern after heat shock.     

Variation in the occurrence of rust (Melampsora spp.) and other diseases and pests, in short-rotation coppice plantations of Salix in the British Isles

A proportional odds model was employed to analyse simple assessments of disease and pest severity in plantings of short-rotation coppice willows at 18 sites in the UK over a 5-year period (1987–1991). Of a wide range of fungal diseases and pests encountered, the most severe and damaging were rusts, caused by Melampsora spp., and feeding damage due to beetles of the family Chrysomelidae. For rust, the model revealed highly significant (P < 0.001) interactions between the main factors, disease development within a season and between seasons, clones and sites. Within seasons, rust severity increased more at sites in the northern than in the southern UK, due mainly to low levels of occurrence at the northern sites early in the season. There was a gradual increase in rust severity over the 5-year period, although clones differed both in terms of disease severity and the extent of change in severity in any one year. Beetle feeding damage was also analysed in a similar way, indicating a general increase from 1987–1989, but also revealing differential patterns over time and between sites.     

Susceptibility of willow clones (Salix spp.) to herbivory by Phyllodecta vulgatissima (L.) and Galerucella lineola (Fab.) (Coleoptera, Chrysomelidae)

Levels of damage by mixed natural infestations of the leaf-feeding chrysomelid beetles, Phyllodecta vulgatissima (L.) (the blue willow beetle) and Galerucella lineola (Fab.) (the brown willow beetle), were determined in replicated field plots of 24 Salix clones at Long Ashton (Bristol, UK) during 1993–94. Over the same period, the host plant preferences of both chrysomelids were investigated in a standard multiple-choice laboratory procedure, where beetles were enclosed in Petri dishes with leaf discs cut from young pot-grown trees propagated from shoot cuttings taken from 20 of the 24 willow clones represented in the field study. The laboratory experiments indicated that P. vulgatissima and G. lineola had similar host plant preferences in the range of willows examined (r >0.85). In both field and laboratory, the least preferred Salix clones and hybrids were those of 5. eriocephala, followed by S. purpurea, S. burjatica, S. dasyclados and S. triandra. Clones of S. eriocephala and S. purpurea were frequently rejected altogether in laboratory tests. Most preferred were clones of S. viminalis and several hybrids of S. viminalis, S. aurita, S. caprea and S. cinerea. These results substantiate the reports that P. vulgatissima and G. lineola are deterred from feeding on willows which have relatively high concentrations of phenolic (salicylate) glucosides in the leaves. The least preferred willows, particularly S. eriocephala, S. purpurea and S. burjatica, could be of great potential value in plant breeding for resistance to these willow beetle pests.     

Assessing the potential of short rotation coppice (SRC) for cleanup of radionuclide-contaminated sites

A small-scale greenhouse investigation was undertaken using Goat willow (Salix caprea) and aspen (Populus tremula) to evaluate the potential of short rotation coppice for remediation of 137Cs- and 90Sr-contaminated sites. Results showed that both species were able to accumulate these radionuclides from a representative disposal soil (aged) and a spiked soil S. caprea accumulating greater levels of 137Cs than P. tremula, with no difference between species for 90Sr accumulation. For each radionuclide, the distribution in both species was similar, with 137Cs accumulation greatest in the roots, whereas 90Sr accumulation was greatest in the leaves. It was also evident that the soil-to-plant transfer factor (Tf) values for 90Sr were greater than for 137Cs, agreeing with differences in the reported bioavailailablity of these radionuclides in soil Based on the Tf values for S. caprea (conservative), estimated remediation times were 92 and 56 yr, for 137Cs and 90Sr, respectively. It is suggested that the selection of Salix species grown in a system of SRC provides a significant opportunity for removal of both 137Cs and 90Sr, primarily due to its higher biomass production. However, for 137Cs phytoremediation investigations into the appropriate use of soil amendments for increasing bioavailability are required.     

Spatio-temporal dynamics of willow beetle (Phratora vulgatissima) in short-rotation coppice willows grown as monocultures or a genetically diverse mixture

1 Phratora vulgatissima (Chrysomelidae) is the major pest of short-rotation coppice willows in the U.K., capable of causing severe defoliation in monoculture plantations. As this beetle shows feeding preferences between willow clones, knowledge of the spatio-temporal dynamics of P. vulgatissima is needed in order to assess the effects of mixed clonal plantings on the management of this pest. 2 Distribution patterns of adult P. vulgatissima were monitored over a season at a spatial scale of every tree or every other tree in single clone plantings (monocultures) of three willow clones and in a regular row mixture of these clones. 3 Distribution indices (deviation from Complete Spatial Randomness) demonstrated that P. vulgatissima adults were spatially aggregated on favourable clones in the monocultures and the mixture. The degree of aggregation (measured by the k-parameter of the negative binomial) differed between willow clones. Beetles were highly aggregated at the start of the season, but less so later. 4 Dispersal studies of P. vulgatissima adults over time showed a pattern of beetle infestation along rows of the preferred clones in a regularly structured mixture. In the monocultures, movement was less directional. 5 Plantation design offers some potential for more effective non-chemical pest management for P. vulgatissima. Monitoring of the pest must take account of our findings that adults occur in mobile aggregations.     

Cadmium and copper uptake and translocation in five willow (Salix L.) species

The efficacy for phytoremediation of five willow species was tested by experimental copper and cadmium uptake in a greenhouse hydroponic system. Five treatments included two concentrations (5 and 25 microM for each metal) and a control. Metal concentrations in solution as well as solution uptake were monitored. Metal resistance was assessed through effects on the dry weight of roots and shoots. The willow species tested were generally resistant of increased Cu and Cd content. Metal accumulation was found in all plant organs of all species. Growth and transpiration were not decreased by 5 microM of copper and 25 microM of cadmium in the solution for most species. 25 microM copper caused injury and reduced the dry weight for all species after 21 d. Salix nigra was highly resistant of both Cu and Cd and accumulated more metals than other species. Future field study should be conducted to confirm the findings and feasibility of the phytoremediation technology using those species.     

Correlation between pathogenicity and molecular characteristics in the willow leaf rusts <Emphasis Type="Italic">Melampsora epitea</Emphasis> and <Emphasis Type="Italic">M. humilis</Emphasis> in Japan

 We studied the correlation between pathogenicity and restriction fragment length polymorphism (RFLP) type, which was determined by polymerase chain reaction-based RFLP analysis of the internal transcribed spacer regions of ribosomal DNA, in the willow leaf rust fungi Melampsora epitea and M. humilis. Eighteen clones of eight Salix species were inoculated with urediniospores from seven collections of the two rust species. M. epitea and M. humilis (RFLP type-5 collections) were pathogenic to six to eight Salix species. RFLP type-7 collections of M. epitea were pathogenic to only two Salix species. The taxonomic relationships of the two rust species are discussed. Received: December 11, 2002 / Accepted: February 17, 2003 RID="*" ID="*" Contribution no. 179, Laboratory of Plant Parasitic Mycology, Institute of Agriculture and Forestry, University of Tsukuba, Japan Acknowledgments We thank K. Katsuya, former professor at the University of Tsukuba, for his encouragement in this study. We are also grateful to M. Yashima, Botanical Garden, University of Tohoku, for his assistance in collecting materials and to R. Suzuki, University of Tsukuba, for providing a rust isolate.     

Nitrate availability and shoot area development in small willow (Salix viminalis)

Abstract. The effect of nitrate supply upon leaf area development in willow ( Salix viminalis ) was investigated. Rooted cuttings were grown in culture solution at two different rates of exponentially-increasing nitrate supply and with free access to other nutrients. During the lag-phase in attaining stable nutrition, the rate of production of unfolded leaves, in the treatment with poorer N-supply, decreased. After acclimation to the different N-availabilities, the rate of dry matter increase per plant nitrogen was constant and the same in both treatments. The ratio of total leaf area to total amount of nitrogen in the plant was slightly higher in the treatment with poorer nitrogen supply. During the exponential (stable) phases of growth, nearly all the increase in total leaf area was attributable to main-stem leaves; large numbers of leaves on axillary shoots in the treatment with better N-supply contributed little to the total area. During the stable growth phase, the rate of production of unfolded leaves was the same in both treatments. Higher maximum values of relative rate of increase in area of single leaves were found with a better N-supply and at higher leaf positions on the main stem. The duration of leaf expansion was not much affected by N-supply. Final leaf size was greater at higher positions on the main-stem. Within a treatment (investigated for poorer N-supply), differences in final leaf size were proportional to differences in numbers of epidermal cells. Final leaf size was greater, at the same node, in the treatment with better N-supply. Better N-availability had increased the final size but not the number of epidermal cells. It was concluded that the higher exponential rate of total area increase with a better N-supply was largely associated with higher rates of expansion in epidermal cells.     

A set of polymorphic SSR loci for subarctic willow (Salix lanata,S. lapponum and S. herbacea)

We report the development of seven microsatellite markers from a library enriched with [CA]₁₅, [GA]₁₅, [AAG]₈ and [ATG]₈. The library was constructed from Icelandic Salix lanata leaf DNA, and seven highly polymorphic single locus products were tested on a range of Scottish and Icelandic S. lanata samples. These seven loci also amplify polymorphic products when tested for cross amplification in a range of Scottish and Icelandic samples of two other subarctic willows (Salix lapponum and Salix herbacea). The microsatellites developed in this study will provide the framework for comparisons with other ecosystem components, and will ultimately inform conservation strategies.     

Correspondence of ectomycorrhizal diversity and colonisation of willows (Salix spp.) grown in short rotation coppice on arable sites and adjacent natural stands

Willows (Salix spp.) are mycorrhizal tree species sometimes cultivated as short rotation coppice (SRC) on arable sites for energy purposes; they are also among the earliest plants colonising primary successional sites in natural stands. The objective of this study was to analyse the degree of colonisation and diversity of ectomycorrhizal (EM) communities on willows grown as SRC in arable soils and their adjacent natural or naturalized stands. Arable sites usually lack ectomycorrhizal host plants before the establishment of SRC, and adjacent natural or naturalized willow stands were hypothesized to be a leading source of ectomycorrhizal inoculum for the SRC. Three test sites including SRC stands (Salix viminalis, Salix dasyclados, and Salix schwerinii) and adjacent natural or naturalized (Salix caprea, Salix fragilis, and Salix?×?mollissima) stands in central Sweden were investigated on EM colonisation and morphotypes, and the fungal partners of 36 of the total 49 EM fungi morphotypes were identified using molecular tools. The frequency of mycorrhizas in the natural/naturalized stands was higher (two sites) or lower (one site) than in the corresponding cultivated stands. Correspondence analysis revealed that some EM taxa (e.g. Agaricales) were mostly associated with cultivated willows, while others (e.g. Thelephorales) were mostly found in natural/naturalized stands. In conclusion, we found strong effects of sites and willow genotype on EM fungi formation, but poor correspondence between the EM fungi abundance and diversity in SRC and their adjacent natural/naturalized stands. The underlying mechanism might be selective promotion of some EM fungi species by more effective spore dispersal.     

Energy requirements and environmental impacts associated with the production of short rotation willow (Salix sp.) chip in Ireland

Willow Salix sp. is currently cultivated as a short rotation forestry crop in Ireland as a source of biomass to contribute to renewable energy goals. The aim of this study is to evaluate the energy requirements and environmental impacts associated with willow (Salix sp.) cultivation, harvest, and transport using life cycle assessment (LCA). In this study, only emissions from the production of the willow chip are included, end‐use emissions from combustion are not considered. In this LCA study, three impact categories are considered; acidification potential, eutrophication potential and global warming potential. In addition, the cumulative energy demand and energy ratio of the system are evaluated. The results identify three key processes in the production chain which contribute most to all impact categories considered; maintenance, harvest and transportation of the crop. Sensitivity analysis on the type of fertilizers used, harvesting technologies and transport distances highlights the effects of these management techniques on overall system performance. Replacement of synthetic fertilizer with biosolids results in a reduction in overall energy demand, but raises acidification potential, eutrophication potential and global warming potential. Rod harvesting compares unfavourably in comparison with direct chip harvesting in each of the impact categories considered due to the additional chipping step required. The results show that dedicated truck transport is preferable to tractor‐trailer transport in terms of energy demand and environmental impacts. Finally, willow chip production compares favourably with coal provision in terms of energy ratio and global warming potential, while achieving a higher energy ratio than peat provision but also a higher global warming potential.     

Patterns of clonal diversity in three species of sub-arctic willow (Salix lanata, Salix lapponum and Salix herbacea)

Sub-arctic willow scrub is an endangered habitat in Britain, and typically occurs on steep crags inaccessible to grazing animals. These willows can reproduce both sexually and asexually, although the relative importance of each is unknown. Knowledge of reproductive mode is important for the design of grazing management and restoration programmes. Accordingly, clonality was assessed in the largest stand of sub-arctic willow scrub in the UK, focusing on Salix lanata and S. lapponum. Little evidence of clonal growth was detected; most individuals possessed distinct multi-locus genotypes. Thus despite the capacity for vegetative reproduction, and seedlings being rarely observed, sexual reproduction is the predominant means of perpetuation and dispersal at this site. We also examined clonal growth in a common willow species (Salix herbacea) that occupies a different habitat type (exposed mountain tops and ridges). Multiple individuals shared identical genotypes up to 7 m apart, suggesting an important role for clonal growth in local patch formation in this species.     

Effects of soil compaction and mechanical damage at harvest on growth and biomass production of short rotation coppice willow

The effects of soil compaction and mechanical damage to stools at harvesting on the growth and biomass production of short rotation coppice (SRC) of willow (Salix viminalis L.) were monitored on clay loam (CL) and sandy loam (SL) soils. Moderate compaction, more typical of current harvesting situations did not reduce biomass yields significantly. Even heavy compaction only reduced stem biomass production by about 12% overall; effects were statistically significant only in the first year of the experiment on sandy loam. Heavy compaction increased soil strength and bulk density down to 0.4 m depth and reduced soil available water and root growth locally. Soil loosening treatments designed to alleviate the effects of heavy compaction did not markedly improve the growth of willow on compacted plots. Hence the focus fell on harvesting. Extensive mechanical damage to stools caused a 9% and 21% reduction in stem dry mass on the clay loam and sandy loam soils as a result of fewer stems being produced. The particularly severe effect on the sandy loam soil probably resulted from a combination of dry conditions in the year of treatment, root damage and soil compaction under stools and might have been aggravated by the young age of the plants (1 year) at the time of treatment.     

Intracellular resistance correlates with initial stage of frost hardening in willow (Salix viminalis)

Distributed model parameters of shoots of five clones of willow ( Salix viminalis ) were examined with electrical impedance analysis at the end of the growing season and with cold acclimation. The parameters were compared with regard to frost hardiness, linoleic (18:2) and linolenic (18:3) fatty acids, unsaturation/saturation ratio of fatty acids and dry weight content. The intracellular resistance (r_i) correlated best with changes in frost hardiness. The value of r_i rose from 1–2 Ω m in non-hardened to about 12 Ω m in hardened samples. In the initial stages of frost hardening, a linear relationship was found between r_i and frost hardiness and levels of 18:2 fatty acid, and an inverse relationship between r_i and levels of 18:3 fatty acid. The unsaturation/saturation ratio of fatty acids rose fairly rapidly in the initial stage of hardening. The dry weight content increased stepwise during the experimental period, and less steadily than r_i. In addition, equivalent circuit parameters changed in the prehardening phase, and were probably connected with cell differentiation and lignification. Frost hardiness by the visual method and by extracellular resistance, determined after controlled freezing tests, correlated well in the initial stages of hardening until about − 10°C but deviated upon further hardening.     

High value of short rotation coppice plantations for phytodiversity in rural landscapes

The demand for wood from short rotation coppice (SRC) plantations as a renewable energy source is currently increasing and could affect biodiversity in agricultural areas. The objective was to evaluate the contribution of SRC plantations to phytodiversity in agricultural landscapes assessed as species richness, species–area relationships, Shannon indices, detrended correspondence analysis on species composition, Sørensen similarities, habitat preference proportions, and species proportions found in only one land use. Vegetation surveys were conducted on 12 willow (Salix spp.) and three poplar (Populus spp.) coppice sites as well as on surrounding arable lands, grasslands and forests in central Sweden and northern Germany. SRC plantations were richer in plant species (mean: 30 species per 100 m²) than arable land (10), coniferous forests (13) and mixed forests in Germany (12). Comparing SRC plantations with other land uses, we found lowest similarities in species composition with arable lands, coniferous forests and German mixed forests and highest similarities with marginal grassland strips, grasslands and Swedish mixed forests. Similarity depended on the SRC tree cover: at increased tree cover, SRC plantations became less similar to grasslands but more similar to forests. The SRC plantations were composed of a mixture of grassland (33%), ruderal (24%) and woodland (15%) species. Species abundance in SRC plantations was more heterogeneous than in arable lands. We conclude that SRC plantations form novel habitats leading to different plant species composition compared to conventional land uses. Their landscape‐scale value for phytodiversity changes depending on harvest cycles and over time. As a structural landscape element, SRC plantations contribute positively to phytodiversity in rural areas, especially in land use mosaics where these plantations are admixed to other land uses with dissimilar plant species composition such as arable land, coniferous forest and, at the German sites, also mixed forest.