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Plant Adaptations to Saturated Soils and the Formation of Hypertrophied Lenticels and Adventitious Roots in Woody Species

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Plant Adaptations to Saturated Soils and the Formation of Hypertrophied Lenticels and Adventitious Roots in Woody Species W&M ScholarWorks Reports 2-1-1996 Plant Adaptations to Saturated Soils and the Formation of Hypertrophied Lenticels and Adventitious Roots in Woody Species Kirk J. Havens Virginia Institute of Marine Science Virginia Institute of Marine Science, Wetlands Program Follow this and additional works at: https://scholarworks.wm.edu/reports Part of the Natural Resources and Conservation Commons Recommended Citation Havens, K. J., & Virginia Institute of Marine Science, Wetlands Program. (1996) Plant Adaptations to Saturated Soils and the Formation of Hypertrophied Lenticels and Adventitious Roots in Woody Species. Wetlands Program Technical Report no. 96-2. Virginia Institute of Marine Science, College of William and Mary. http://dx.doi.org/doi:10.21220/m2-26g9-p109 This Report is brought to you for free and open access by W&M ScholarWorks. It has been accepted for inclusion in Reports by an authorized administrator of W&M ScholarWorks. For more information, please contact [email protected]. Plant Adaptations to Saturated Soils and the Formation of Hypertrophied Lenticels and Adventitious Roots in Woody Species Kirk J. Havens Determining the extent of forested wetlands is a (1920) definition of a subgroup called helophytes difficult task. These areas are subject to overflow (Greek helos = marsh) may be more appropriate and inundation of varying frequenciesand dura­ for species that can.occur equally in both wetland tions. The frequency, timing, and duration of and upland conditions. Clements (1920) defined inundation and soil saturation are the driving helophytes as "amphibious plants rooted in forces behind the processes that affect species water or mud." composition. Identifying that portion of a bot­ tomland hardwood forest that is a wetland With the advent of laws regulating the develop­ within the meaning of Section 404 of the Federal ment of wetlands in the United States, a series of Water Pollution Control Act and which meets the additional terms defining plant adaptation to wet definition of wetlands as determined by the regu­ conditions was introduced. This action effec­ lating agency, U.S. Army Corps of Engineers, has tively created five groupings of plants based on been a controversial issue. The presence of cer­ differences in the probability of occurrence in tain adaptations that trees undergo to cope with wetland versus upland habitats. These groups flooding can provide information on the dura­ are termed: obligate upland, facultative upland, tion of flooding at a specific site. This could be facultative, facultative wetland, obligate wet­ valuable to resource managers by helping in the land. Facultative plants are defined as having an determination of the extent of forested wetland equal probability of occurring in wetland or up­ systems. land habitat. The probability of occurrence in wetlands of plants in each grouping is listed in A number of plant species are known to grow in Table 1 (see page 2). soil conditions that vary from dry upland to per­ manently inundated wetlands. Wetlands can be The adaptive abilities of plants allow them to considered transitional areas that occur along a survive in a broad range of physical conditions. soil-moisture gradient between uplands and Even though a facultative species can be found shallow water habitat. Hydrologic regimes asso­ growing in upland, dry conditions, it may pos­ ciated with wetlands range from permanent in­ sess physiological or morphological adaptations undation by shallow water to periodic soil that also allow it to survive in wetlands. saturation (Tiner, 1991). The flooding of soil causes a relatively rapid de­ Plants adapted to living in conditions of periodic pletion of oxygen. Oxygen can be depleted from saturationor inundation are called hydrophytes the soil in a few hours to a few days due to (Greek hydro = water). However, Clements' continued root oxygen demand, microbial respi- ration, and chemical oxidation of reduced Wetland Probability of organic and inorganic compounds. Dis- 12 2 Indicator Category Occurrence in Wetlands solved oxygen d1.ffu s1on, . at 1 x 10- g cm- sec-1 into the waterlogged soil, is 10,000 Obligate Wetland (OBL) >99% times slower than in non-waterlogged soil Facultative Wetland (FACW) 67-99% (Armstrong, 1967) and 100 times slower Facultative (FAC) 34-66% than oxygen consumption (Ponnampe­ Facultative Upland (FACU) 1-33% ruma, 1972). After the loss of oxygen from Obligate Upland (UPL) <1% the system, microbes begin using oxidized soil components in their respiration which Table J. Wetland indicator categories of plant species under lowers the redox potential of the system. natural conditions (adapted from Reed, l 988). Redox potential (Eh) is a measure of the electron Patrick and Mahapatra (1968) classified soil re­ availability in chemical and biological systems. dox conditions into four ranges at pH 7: oxidized Electrons are essential reactants in inorganic, or­ soils 400 m V; moderately reduced soils 400 to 100 ganic, and biochemical reactions. Chemical spe­ mV; reduced soils 100 to -100 mV; and highly cies that lose electrons become oxidized while reduced soils -100 to -300 m V. Redox potentials those that gain electrons become reduced. In for several reactions in soils of pH 7 from Bohn aqueous solutions, redox potential affects the oxi­ (1971) and Ponnamperuma (1984) are shown in dation states of oxygen, hydrogen, nitrogen, Table 2. manganese, iron, copper, cobalt, sulfur, and carb­ on in succession (Bohn, 1971). As each of these As the system becomes more reduced, plants elements is reduced it indicates the intensity of must cope with the increased concentrations of oxidation or reduction of a chemical or biological phytotoxins such as iron, manganese, and sul­ system. phide (Ernst, 1990) and the increased stress of being rooted in a waterlogged, anaerobic (with­ out oxygen) environment. Wetlands Program School of Marine Science Wetland plants have a number of responses and Virginia Institute of Marine Science adaptations such as the formation of adventitious College of William and Mary water roots and hypertrophied lenticels that al­ Gloucester Point, Virginia 23062 low them to cope with the effects of waterlogging (Figure 1). Hypertrophied lenticels are enlarged Dr. Carl Hershner, Program Director cells that allow for increased oxygen diffusion Publisher by: VIMS Publication Center along the stem. Adventitious roots are porous roots which are more capable of oxygen uptake This technical report was funded, in part, by the Department of Environmental and generally form off the primary root or off the Quality's Coastal Resources Management stem near the water line. One particularly impor­ Program through Grant #NA470Z0287-01 tant adaptation is the ability to transport oxygen of the National Oceanic and Atmospheric from the stem to the roots via aerenchyma tissue Administration, Office of Ocean and Coastal Resources (Figure 2.., page 4) and the subsequent oxidation Management, under the Coastal Zone Management Act of of the root zone (rhizosphere) by radial loss of 1972, as amended. oxygen from the roots. Commonwealth's Declared Policy: Oxygen enters the xylem of the stem of trees "to preserve the wetlands and to prevent their through the leaves and lenticels and hypertro­ despoliation and destruction ... " phied lenticels at the waterline are considered the main entry point (Armstrong, 1978; Coutts and Armstrong, 1971). Blockage of the hypertro- Printed on recycled paper. Ci) 2 Hook and others (1971), Bohn (1971) Ponnamperuma '1984) and Scholander and others (1955) have demonstrated +225 mV Nitrate reauction +330 m V Disappearance of d the importance of lenticels +200 mV Manganese reduction +220 m V Disappearance of NCn­ 2 for aeration in willows, tu­ +120 m V Iron reduction +200 m V AppearanceofMn + 2 pelos, and mangroves. In a +100 m V Sulfate reduction +120 m V Appearance of Fe + classic experiment on man­ -150 m V Disappearance of soi- groves, it was demon­ -250 m V Appearance of Cfu strated that the oxygen concentration in roots sub­ Table 2. Redox potentials in soils of pH 7. merged in anaerobic mud remained continuously phied lenticels located at the waterline has been high at 15-18%, but if the hypertrophied lenticels shown to reduce the oxygen status of the roots were blocked the concentration fell to 2% or less (Armstrong, 1968; Hook et al., 1971; Scholander (Scholander et al., 1955). et al., 1955). Due to the resistance to the move­ ment of oxygen and its consumption by respira­ Adenosine triphosphate (ATP) is the major en­ tion as it moves through the stem to the root, it is ergy-carrying molecule in cellular metabolism believed that oxygen supplied through leaf en­ and the generation of ATP is much more efficient trance is of little importance to the roots of trees under aerobic conditions. Without some mecha­ (Coutts and Armstrong, 1971). Armstrong (1968), nism for ATP generation during anaerobic condi- Adventitious root extending from hypertrophied lenticel Hypertrophied lenticels Figure I . Hypertrophied lenticels and adventitious roots on the stem of a red maple seedling. 3 tions, the amount of ATP in a root cell would be rhizosphere, neutralizing phytotoxins and sup­ sufficient for less than one minute of metabolism plying oxygen to rhizosphere microbial popula­ (Roberts et al., 1984). The diffusion of oxygen to tions. In addition, the iron oxide plaque that the roots allows for. continued root growth and forms along the roots can scavenge other metallic nutrient uptake under anaerobic conditions. Ra­ ions thereby decreasing their toxic effects (Crow­ dial leakage of oxygen from the root oxidizes the der, 1991). Waterlogging causes an in­ crease in ethylene production in tissues below and immedi­ ately above the water line (Yang, 1980; Tang and Ko­ zlowski, 1_984) resulting in cel­ lulase activity. Cellulase softens the cell walls causing stem hypertrophy (enlarge­ ment) and development of aer­ enchyma tissue in both the lenticels and the adventitious roots (Sena Gomes and Ko­ zlowski, 1980; Kawase, 1981; Kozlowski, 1984). Porosity is increased due to the formation of large aerenchyma cells and the large channels in the root cortex enhance oxygen trans­ port (Justin and Armstrong, 1987).
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