To determine the tolerance of Salix gracilistyla to repetitive alternate flooding and drought, we measured leaf stomatal conductance, pre-dawn water potential, osmotic adjustment, and biomass production under greenhouse conditions. We used a control and nine crossed treatments (F1-D1-F3-D3) in which we combined 1-, 2-, or 3-week floodings (F) and droughts (D). Leaf stomatal conductance was lowest in 3 weeks of flooding or drought when the preceding event (flood or drought) was also of a 3-week duration. Leaf pre-dawn water potential was reduced in 3 weeks of drought when preceded by 2 or 3 weeks of flooding. Cuttings had slight osmotic adjustments in repetitions of long floodings and droughts. During longer durations of drought in crossed experiments, plants had low root and shoot mass, few hypertrophic lenticels, and reduced leaf mass; when flooding duration increased in crossed experiments, root mass was reduced, there were more hypertrophic lenticels, and the leaf area was reduced. Cuttings achieved stress tolerance by inhibition of transpiration, osmotic adjustment, reduction of transpiration area, and development of hypertrophic lenticels. Stress tolerance was weak when repetitive 2-or 3-week floodings were combined with 3-week droughts. The duration of flooding and drought periods under which S. gracilistyla achieves stress tolerance may be critical in determining distributions along riverbanks.
The objective of this study is to determine the effects of substrate moisture and oxygen availability on growth traits of Salix gracilistyla Miquel, which colonizes gravel bars along rivers, the shoot growth schedule, biomass production, and resource allocation were examined under greenhouse conditions. We used four treatments representing a range of substrate moisture and oxygen availability: drought (D), flooding with standing water (FS), flooding with running water (FR), and control without drought or flooding (C). Cuttings in D stopped flushing and had low biomass production, reduced total leaf mass, and small leaves. Under anaerobic conditions, cuttings in FS stopped flushing and had low biomass production, small root biomass, low biomass allocation to roots, shallow roots, high biomass allocation to hypertrophied lenticels, and a few small, thick leaves. Under aerobic conditions, cuttings in FR showed continuous branch elongation and flushing, large biomass production, and large leaf biomass, similar to cuttings in C, in addition to low allocation to hypertrophied lenticels and many large leaves. The growth of cuttings was not inhibited by flooding of the roots throughout the experiment unless the conditions were anaerobic. Thus, cuttings respond to water stress under low moisture conditions by reducing the transpiration area and respond to flooding under low oxygen conditions by high allocation to hypertrophied lenticels and reduced transpiration area. Plasticity in the shoot growth schedule, biomass production, and resource allocation according to moisture conditions and the ability to develop hypertrophied lenticels upon flooding allow S. gracilistyla to colonize sites in which both desiccation and flooding occur.
To clarify the effects of inundation on the establishment of Salix gracilistyla trees, the relationship between total inundation duration and the distribution and size of trees was examined on a gravel bar along the Miya River in central Japan. The density of S. gracilistyla was low at both low and high elevations on the gravel bar, but high at middle elevations. The density increased with up to 165 days of inundation and declined gradually with more than 165 days of inundation. Density had a negative effect on crown area, whereas inundation duration had a positive effect. At lower elevations, growth limitation of S. gracilistyla is mainly caused by inundation stress, whereas at higher elevations it seems to be primarily due to drought stress, although it depends on the total duration of inundation in a year (TDIY). Consequently, the density is highest at middle elevations where trees do not become too large. These results suggest that inundation duration is one factor determining the density of S. gracilistyla trees, which in turn affects the size of each tree.
To compare the responses to repeated flooding and drought of Salix gracilistyla, which grows on coarse gravel substrates, and Salix subfragilis, which grows on fine silt or clay substrates, we measured pre-dawn leaf water potential (Ww pd ), osmotic adjustment (Ww tlp , Wo sat ), and biomass production of cuttings under greenhouse conditions. The experimental design involved a control and four treatments that crossed 1 or 3-week flooding (F) with 1 or 2-week droughts (D). Ww pd was reduced after 2 weeks of drought when preceded by 1 week of flooding. Neither species increased osmotic adjustment in response to increased duration of drought between repeated 3-week flooding. Moreover, a decrease in the ratio of leaf biomass to total biomass or an increase in the ratio of root biomass to total biomass with longer drought repetitions was not observed for either species. The root ratio of S. gracilistyla was more strongly inhibited by flooding than that of S. subfragilis. The shoot-to-root ratio of S. subfragilis was higher than that of S. gracilistyla in all F combinations. The hypertrophied lenticel ratio of S. gracilistyla after 1 week of flooding was nearly the same as that after 3 weeks of flooding, whereas values for S. subfragilis after 1 week of flooding were lower than those after 3 weeks of flooding. The low allocation to roots and the generation of hypertrophied lenticels by S. gracilistyla in response to flooding, as compared with S. subfragilis, seem to be related to the different habitat substrate conditions of the two species.
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