Population Differentiation along a Flood Frequency Gradient: Physiological Adaptations to Flooding in Nyssa sylvatica

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Abstract
Throughout the southeastern United States the hardwood Nyssa sylvatica (sensu lato) is distributed along a soil moisture gradient from upland sites, which are never flooded, to floodplains, which are periodically flooded and drained to permanently flooded swamps. Population differentiation with respect to flood tolerance and related physiological attributes was investigated using 1—year—old seedlings grown in a greenhouse from seed collected along this gradient. Upland plants were very intolerant of flooded soils. Their root systems deteriorated, root respiration rates dropped and, after a year under such conditions, survival was poor and those that did remain were greatly stunted and had accumulated large concentrations of many nutrient elements. In contrast swamp plants were quite tolerant of flooded soils. Upon flooding, parts of the orginal root system were lost but new roots were initiated which had an increased capacity for alcoholic fermentation. Many of these new roots were more succulent, larger in diameter, and less branched than drained roots. Such succulent roots however were only a temporary response to short—term flooding; plants flooded for a year did not have such roots, rather the root system superficially resembled drained roots. Concomitant with this return to drained—like roots was an increase in internal oxygen transport to the roots and a drop in alcoholic fermentation. Floodplain plants under drained conditions allocated less biomass to roots than to shoots and had high respiration rates, traits similar to upland plants. Under flooded conditions they initiated new roots with medium respiration rates, allocated less biomass to roots than to shoots, significantly increased oxygen transport to the roots and had high survival, traits similar to swamp plants. Thus, the floodplain population produced a distinctly flood—tolerant phenotype; but not nearly as tolerant of flooded conditions as the swamp phenotype. Floodplain plants differed from swamp plants in transporting less oxygen to the roots under drained conditions, initiating fewer succulent—type roots and not accelerating alcoholic fermentation upon flooding and after a year under flooded conditions having less total biomass, less oxygen transport to the roots and a greater accumulation of Fe and Mn in the roots. The floodplain plants apparently have been selected to be similar to upland plants under drained conditions and swamp plants under flooding and one consequence of this is that their tolerance of flooded conditions is intermediate. It is argued that one of the more important trade—offs in adapting to flooded conditions is that high internal oxygen transport carries with it a ‘cost’ in terms of excessive water loss under water stress conditions.