Martin Weih

Characterising willows for biomass and phytoremediation: growth, nitrogen and water use of 14 willow clones under different irrigation and fertilisation regimes

Fourteen clones of willow (Salix spp.) were characterised in terms of growth, nitrogen and water-use efficiency under different irrigation and fertilisation treatments. Cuttings of willow clones, some commercially introduced and others new material, were pot-grown outdoors in Central Sweden under four experimental treatments in a full-factorial design. The experiment covered the period from bud-break until leaf abscission and the experimental conditions included two irrigation and two fertilisation regimes. The growth of the clones was evaluated in terms of relative growth rate and total biomass production of whole plants and shoots. Nitrogen (N) economy was studied by means of N productivity, N accumulation and N losses by leaf abscission. Water economy was analysed with respect to intrinsic water-use efficiency (foliar carbon isotope ratio; δ13C) and the capacity of leaves to retain water (relative water content). Significant differences between clones were found in nearly all parameters measured and the clones varied in the responses to the experimental treatments (clone × factor interaction effects). Thus, clone ranking often changed depending on the experimental treatment. The results are discussed with respect to clone selection for different willow applications such as biomass production and phytoremediation, and willow growth performance under different water and nutrient availabilities. The growth-physiological characterisation of young willows in the short term (several months) is regarded as a suitable approach for pre-selection of promising clones prior to extensive field evaluation.

Short-rotation forestry with hybrid aspen (Populus tremula L.×P. tremuloides Michx.) in Northern Europe

Abstract This review summarises the knowledge about the ecology, breeding and management of hybrid aspen (Populus×wettsteinii Hämet-Ahti=P. tremula L.×P. tremuloides Michx.). The review is restricted mainly to Northern Europe, where hybrid aspen has been most intensively studied and cultivated and where it has proved to be one of the fastest-growing hardwoods, suitable for the production of pulp- and energy-wood using the principles of short-rotation forestry. During recent decades over 4500 ha have been cultivated with hybrid aspen in the region. A number of research articles and domestic project reports involving hybrid aspen have been published, providing the basis for this review. Breeding has resulted in clones with high productivity and improved resistance to pests and diseases. Thus, hybrid aspen has fulfilled the preconditions for becoming an economically valuable hardwood in Northern Europe. Hybrid aspen plantations can be established on abandoned agricultural land, on forest land, and for the reclamation of exhausted surface mines. However, fast growth rate occurs only in fertile sites with good nutritional and hydrophysical properties. An increased area of Populus plantations on forest or agricultural land can have both positive and negative impacts on biodiversity, depending on landscape context, management activities and considered organisms. Further studies are needed concerning silviculture, site-growth relations, stability of clones, environmental and biodiversity impacts in large-scale plantations at various sites and adaptation of hybrid aspen to climate change.

Plant stoichiometry at different scales: element concentration patterns reflect environment more than genotype

All plant species require at least 16 elements for their growth and survival but the relative requirements and the variability at different organizational scales is not well understood. We use a fertiliser experiment with six willow (Salix spp.) genotypes to evaluate a methodology based on Euclidian distances for stoichiometric analysis of the variability in leaf nutrient relations of twelve of those (C, N, P, K, Ca, Mg, Mn, S, Fe, Zn, B, Cu) plus Na and Al. Differences in availability of the elements in the environment was the major driver of variation. Variability between leaves within a plant or between individuals of the same genotype growing in close proximity was as large as variability between genotypes. Elements could be grouped by influence on growth: N, P, S and Mn concentrations follow each other and increase with growth rate; K, Ca and Mg uptake follow the increase in biomass; but uptake of Fe, B, Zn and Al seems to be limited. The position of Cu lies between the first two groups. Only for Na is there a difference in element concentrations between genotypes. The three groups of elements can be associated with different biochemical functions.

Willow genotype, but not drought treatment, affects foliar phenolic concentrations and leaf-beetle resistance

In a greenhouse experiment we examined the effect of willow genotype and irrigation regime (moderate drought and well‐watered) on plant growth parameters, foliar nitrogen, and phenolic concentrations, as well as on the preference and performance of the blue leaf beetle, Phratora vulgatissima (L.) (Coleoptera: Chrysomelidae). The 10 vegetatively propagated willow genotypes in the experiments were F2 full‐sibling hybrids, originated from a cross between Salix viminalis (L.) (Salicaceae) (high in condensed tannins) and Salix dasyclados (L.) (Salicaceae) (rich in phenolic glycosides). Insect bioassays were conducted on detached leaves in Petri dishes as well as with free‐living insects on intact potted plants. The 10‐week long irrigation treatments caused statistically significant phenotypic differences in the potted willow saplings. Total biomass was somewhat higher in the well‐watered treatment. The root to total biomass ratio was higher in the drought‐treatment plants. There was significant genotypic variation in foliar nitrogen concentrations, and they were higher in the drought‐treatment plants. There was also a strong genotypic variation in each of the phenolic substances analyzed. Condensed tannins, which accounted for the greatest proportion of total phenolic mass, were higher in the well‐watered treatment. There was, however, no difference in levels of the other phenolics (salicylates, cinnamic acid, flavonoids, and chlorogenic acid) between irrigation treatments. The sum of these phenolics was higher in the well‐watered treatment. There was a strong variation in P. vulgatissima larval development on different willow genotypes, and larval performance was negatively correlated with levels of salicylates and cinnamic acid. There was, however, no effect of irrigation treatment on larval performance. Phratora vulgatissima preferred to feed on well‐watered plants, and we found a preference for oviposition there, but neither feeding nor oviposition site preference was affected by willow genotype. Adult feeding and oviposition preferences were not correlated with larval performance.

QTL analyses of drought tolerance and growth for a Salix dasyclados × Salix viminalis hybrid in contrasting water regimes

Quantitative trait loci (QTL) for growth traits and water-use efficiency have been identified in two water regimes (normal and drought-treated) and for a treatment index. A tetraploid hybrid F2 population originating from a cross between a Salix dasyclados clone (SW901290) and a Salix viminalis clone (‘Jorunn’) was used in the study. The growth response of each individual including both above and below ground dry-matter production (i.e. shoot length, shoot diameter, aboveground and root dry weight, internode length, root dry weight/total dry weight, relative growth rate and leaf nitrogen content) was analysed in a replicated block experiment with two water treatments. A composite interval mapping approach was used to estimate number of QTL, the magnitude of the QTL and their position on genetic linkage maps. QTL specific for each treatment and for the treatment index were found, but QTL common across the treatments and the treatment index were also detected. Each QTL explained from 8% to 29% of the phenotypic variation, depending on trait and treatment. Clusters of QTL for different traits were mapped close to each other at several linkage groups, indicating either a common genetic base or tightly linked QTL. Common QTL identified between treatments and treatment index in the complex trait dry weight can be useful tools in the breeding and selection for drought stress tolerance in Salix.

Above-ground Woody Biomass Production of Short-rotation Populus Plantations on Agricultural Land in Sweden

Although poplars are widely grown in short-rotation forestry in many countries, little is known about poplar growth performance in Sweden. In this study, above-ground biomass production was estimated for several hybrid aspen and poplar clones planted at different initial density at five locations across Sweden. Biomass assessments were based on allometric relationships between total above-ground woody dry weight and the diameter at breast height. According to a common harvest practice, tree biomass was partitioned into pulpwood and biomass for energy purposes. The percentage of pulpwood was strongly determined by clone for DBH >10 cm. The mean annual increment ranged from 3.3 Mg ha−1 yr−1 for balsam poplar in the north to 9.2 Mg ha−1 yr−1 for 9-yr-old ‘Boelare’ in southern Sweden. At the same age, hybrid aspen reached 7.9 Mg ha−1 yr−1. The results suggest that poplars and hybrid aspen are superior as biomass producers compared with tree species commonly grown on agricultural land at these latitudes. The results are discussed in the light of future wood supply for pulpwood and energy purposes in Sweden.

Influence of young poplar stands on floristic diversity in agricultural landscapes (Sweden)

Abstract Floristic diversity in 21 small short-rotation poplar plantations, 0.1 to 13 ha in size and aged 6 to 14 yr, was compared with adjacent agricultural fields of various crop types. The poplars were grown on former agricultural land and most of the study sites were located in southern and central Sweden. Surveys of ground vegetation were carried out by using nested quadrats (plots of 0.25 and 0.016 m 2 size) located along normally five transects running perpendicular to the habitat boundaries. Mean cumulative species number across all sites (all transects and plots included) was around 20 and similar in poplar stands and arable fields. However, the two habitat types shared only 3 to 12 species (mean: 8 species). Mean species number per survey plot was 4.5 in poplar stands compared to 5.3 in arable fields. Classification (TWINSPAN) and Detrended Correspondence Analysis (DCA) showed some floristic separation of the two habitat types. The results were interpreted in a landscape context. Mainly based on the low quantity of species shared by poplar stands and adjacent arable fields, the results support the hypothesis that small-scale poplar plantations increase floristic diversity in landscapes dominated by agriculture. Compared to old-growth mixed deciduous forests in different regions of Sweden, species richness of poplar plantations was similar or lower. Die floristische Artenvielfalt in 21 kleinflachigen Pappelbestanden (0,1–13 ha) im Kurzumtrieb wurde mit derjenigen benachbarter cker verglichen. Die Pappeln waren 6–14 Jahre alt und wuchsen auf ehemaligen Ackerflachen, vorwiegend in Sud- und Mittelschweden (ein Standort in Nordschweden). Die Bodenvegetation wurde mit Hilfe ineinander geschachtelter Aufnahmequadrate erfasst (Aufnahmeflachen 0,25 und 0,016 m 2 ). In jedem Bestand und Feld wurden funf parallel zueinander liegende Transekte angelegt, die senkrecht vom Bestandesrand ins Bestandesinnere verliefen. In jedem Transekt lagen in definierten Abstanden zum Bestandesrand funf Aufnahmeflachen. Die mittlere Gesamtartenzahl uber alle Bestande war 20 und etwa gleich in Pappelbestanden und ckern. Allerdings waren nur zwischen 3 und 12 (Mittelwert: 8) Arten gemeinsam in Pappelbestanden und benachbarten ckern. Die Artenanzahl pro Aufnahmeflache war kleiner in Pappelbestanden als in ckern. Klassifikation (TWINSPAN) und Ordination (DCA) zeigte eine gewisse Trennung der beiden Vegetationstypen. Die Ergebnisse wurden auf Landschaftsebene interpretiert und stutzen die Hypothese, dass kleinflachige Pappelplantagen die floristische Artenvielfalt v.a. in intensiv genutzter Agrarlandschaft erhohen kann. Im Vergleich zu naturlichen Laubmischwaldern verschiedener Regionen Schwedens war die Artenvielfalt in den Pappelplantagen entweder ahnlich (ostliches Mittelschweden) oder niedriger (Sudostschweden).

Assessment of nutrient use in annual and perennial crops: A functional concept for analyzing nitrogen use efficiency

The use of more nutrient-efficient crops is important for maintaining yields while enhancing environmental sustainability. Various approaches are being applied to evaluate aspects of plant nutrient use efficiency, among them ecological concepts based on accumulation and losses of biomass and nutrients, agronomic concepts with a major focus on agricultural crops and harvested products, and physiological approaches assessing single physiological processes important for nutrient use. Unfortunately, the various approaches are often not compatible. Here we propose, with the example of nitrogen (N) use efficiency (NUE) of cereals, to integrate the functionally important components of NUE in a common conceptual framework. We link productivity to N in crops and seeds and consider the whole life-cycle of the crop (including seeds). Three major components of NUE are separated: The N uptake efficiency, grain-specific N efficiency and grain N concentration. The three components combine to a measure of overall NUE in terms of the N yield in harvested grain per unit of N in seed grain or soil N. The concept can be applied for both annual and perennial plants, which is demonstrated with the examples of winter wheat and a perennial energy crop (Salix) grown in Central Sweden.

Genetic and environmental variation in spring and autumn phenology of biomass willows (Salix spp.): effects on shoot growth and nitrogen economy

Six commercial willow (Salix spp.) varieties were examined to investigate the effects of genotype and environment on spring and autumn phenology and the relationships between phenology, shoot growth and leaf nitrogen (N) retranslocation. The willows were field-grown under different irrigation and fertilization in central Sweden. Two independent data sets of bud-burst, leaf unfolding duration, growth cessation and the timing of leaf abscission were assessed, and the biomass and leaf N data from the end of the first cutting cycle were used. Specific hypotheses were that (1) spring phenology has a greater effect on the shoot biomass production than autumn phenology; (2) later bud-burst is associated with more rapid leaf unfolding; (3) the timing of leaf abscission has a greater effect on the shoot biomass production than height growth cessation; and (4) later leaf fall is associated with poorer leaf N retranslocation. Bud-burst date varied by 19 and 39 days in the 2 years and leaf unfolding duration varied by 13 and 38 days. Growth cessation varied by 2.5 weeks and completion of leaf abscission (> 90% of leaves shed) by more than 3 weeks between the genotypes and treatments. Bud-burst date was inversely correlated with leaf unfolding duration (R(2) = 0.96). Significant effects of the duration of leafy period (bud-burst to leaf abscission) and bud-burst date on shoot growth were found. Delayed growth cessation and leaf abscission were generally associated with a greater biomass production, but especially the relationship between growth cessation and biomass was weak. The results show that the timing of bud-burst and leaf abscission is more important for willow biomass production than growth cessation. Delayed leaf abscission has a negative effect on leaf N retranslocation and increases the N losses. The results have implications for the breeding of perennial energy crops.

Optimizing nitrogen economy under drought: increased leaf nitrogen is an acclimation to water stress in willow (Salix spp.)

Background and Aims The major objective was to identify plant traits functionally important for optimization of shoot growth and nitrogen (N) economy under drought. Although increased leaf N content (area basis) has been observed in dry environments and theory predicts increased leaf N to be an acclimation to drought, experimental evidence for the prediction is rare. Methods A pedigree of 200 full-sibling hybrid willows was pot-grown in a glasshouse in three replicate blocks and exposed to two water regimes for 3 weeks. Drought conditions were simulated as repeated periods of water shortage. The total leaf mass and area, leaf area efficiency (shoot growth per unit leaf area, EA), area-based leaf N content (NA), total leaf N pool (NL) and leaf N efficiency (shoot growth per unit leaf N, EN) were assessed. Key Results In the water-stress treatment, shoot biomass growth was N limited in the genotypes with low NL, but increasingly limited by other factors in the genotypes with greatest NL. The NA was increased by drought, and drought-induced shift in NA varied between genotypes (significant G × E). Judged from the EA–NA relationship, optimal NA was 16 % higher in the water-stress compared with the well-watered treatment. Biomass allocation to leaves and shoots varied between treatments, but the treatment response of the leaf : shoot ratio was similar across all genotypes. Conclusions It is concluded that N-uptake efficiency and leaf N efficiency are important traits to improve growth under drought. Increased leaf N content (area basis) is an acclimation to optimize N economy under drought. The leaf N content is an interesting trait for breeding of willow bioenergy crops in a climate change future. In contrast, leaf biomass allocation is a less interesting breeding target to improve yield under drought.