Biological Adaptations to the Wetland Environment Two problems that biota face in wetlands 1. Anoxia - depletion of oxygen in soil and water in a flooded wetland 2. Salt - changes osmotic potential and when concentrations in the cell are high, it can be toxic (inactivate metabolic enzymes) Bacteria Cyanobacteria Single-celled protists Adaptations to anoxia -anaerobic respiration ex. Thiobacillus and Desulfovibrio are bacteria that are part of sulfur cycle. Thiobacillus oxidixes elemental sulfur (or hydrogen sulfide) to form sulfates. Desulfovibrio uses sulfates as the terminal electron acceptor and hydrogen sulfide is produced Adaptations to salt -active transport using sodium potassium pumps to accumulate potassium and dilute sodium Plants-Mosses Sphagnum or peat moss Liverwort Ferns Horsetails Gymnosperm Angiosperm Adaptations to anoxia -anaerobic respiration Plants that are adapted to wetlands contain enzymes other than ADH and are not poisoned by alcohol accumulation. MDH that produces malate is a common example of an enzyme utilized for anaerobic respiration in wetland plants. Aerenchyma and adventitious root development above anaerobic zone are stimulated by hormone auxin that triggers growth of adventitious roots. New growth releases ethylene that accumulates in roots under anaerobic conditions such as waterlogged soils. Ethylene stimulates the formation ofcellulase that opens up cortex as aerenchyma. Examples of common plants with adventitous roots and aerenchyma are water lilies, mangroves, and bald cypress. Water lilies have floating leaves. Mangroves and bald cypresses have pneumatophores that contain aerenchyma and function as respiratory organs. Buttress roots often contain aerenchyma and provide support for woody plants. hypertrophied lenticels - enlarged lenticels that are more numerous on stem increase oxygen uptake during inundation rhizosphere oxidation - oxygen diffusing from the roots reacts with potentially toxic reduced ions forming oxidized forms that precipitate out of the rhizosphere (ie iron(II) to iron (III) and manganese (II) to manganese (III)). Adaptations to salt 1) periderm, a thickened epidermis, is a barrier protecting inner tissues. 2) salt secreting glands - secretory organs on leaves. 3) C-4 photosynthesis reduces water loss via transpiration when stomates are closed during dry or saline conditions. C-3 photosynthesis works well as long as stomates remain open. C-4 photosynthesis keeps carbon dioxide concentration high so photorespiration does not occur. Photorespiration when stomates Close produces a 2-C compound that cannot be used to make sugar. Adaptations involving nutrient absorption 1) water - plants intolerant to anaerobic environments have poor root metabolism and respond as if in drought situation- stomates close, decreased transpiration, and wilting. Countered by adventitous roots, aerenchyma, and enlarged lenticels 2) nitrogen - In wetlands, nitrates are reduced to ammonium that is toxic. Wetland plants maintain normal rates of nitrogen uptake by converting ammonium to nitrate in rhizosphere or have the ability to absorb ammonium directly and convert it to amides. Others have increased levels of nitrate reductase so that when nitrates are absorbed they can more efficiently convert them to amide groups to be used in synthesizing amino acids. Some wetland plants are carnivorous and obtain nitrogen from exoskeletons. 3) phosphorous - availability increases in waterlogged soils. 4) iron and manganese - increase can be toxic by inactivating enzymes. Rhizosphere reduces that toxicity by oxidation and subsequent precipitation. 5) sulfur - sulfides produced by anaerobic bacteria are toxic. They affect enzymes such as those needed for respiration. May be overcome by oxidation in rhizosphere and accumulation of sulfates in vacuoles. Adaptations involving reproduction 1) timing of seed production innonflood season by delayed flowering 2) hydrochory. Fruits or seeds adapted to floating and fungal resistant. 3) vivipary. Seeds germinate on the plants and seedlings drop into the water. Example: mangroves 4) prolonged seed viability Ex bald cypress seeds remain viable for 20 years or more and germinate during infrequent periods when soil isdewatered. 5) flood tolerant seeds and seedlings. Some species can germinate underwater or have seedlings that survive inundation. For example ashes have seedlings that can survive inundation. Animals have six major adaptations to anoxia. 1) Modifications for gaseous exchange. Examples include gills andparapodia. 2) Mechanisms to improve the oxygen gradient across a membrane. For example, cilia moving water over gills or animals swimming to oxygenrich water. 3) Better circulatory system. 4) Use of more efficient respiratory pigments in blood and/or higher concentrations of these pigments. For example, mollusks have hemocyanin (copper based) while the annelids have more efficient hemoglobin. 5) Low oxygen behaviors such as reduced motility or closing a shell. 6) Increased metabolism or heart pumping rates. Three Major Adaptations to Salt in Animals 1) nephridia help retain water and eliminate salts in concentrated urines. 2) salt secreting glands. 3) Reproduction with large numbers of larvae that are distributed widely and can tolerate a variety of salinity changes so that some survive. Plants are wetland indicators because of their adaptations and demarcate wetland boundaries. Some common angiosperm plant families and a few examples of species found in North-central Texas Wetlands Aceraceae (maple) Trees or shrubs with reduced flowers that produce winged fruits called samaras. Morphological adaptations: adventitious roots, hypertrophied lenticels. Acer negundo Alismataceae (arrowhead) Herbs with linear to lanceolate or arrowhead-shaped leaves with three sepals and petals as well as many stamens and pistils. Most are in wetland habitats. Morphological adaptations: polymorphic leaves (linear when submerged, broader when emergent). Asteraceae (sunflower) Small flowers in heads. Physiological adaptation: Senecio vulgaris has a slight increase in metabolism under anaerobic conditions as compared to non-wetland species that have large increases. Cupressaceae (cypress or cedar) Trees or shrubs with scale-like leaves and small woody cones. Morphological adaptations: buttressed trunks and pneumatophores. Cyperaceae (sedge) Grasslike herbs often having edged stems and a single bract covering each of its many reduced flowers. Most are in wetland habitats. Morphological adaptations: inflated stems and leaves Physiological adaptations: Some have been shown to have malate accumulation and lack of ADH activity. Cyperus strigosus Carex Juncaceae (rush) Grasslike herbs with round stems and basal, tufted leaves as well as six papery flower parts. Most are in wetland habitats. Morphological adaptations: inflated stems and leaves Physiological adaptation: Juncus effusus can grow roots under very low oxygen conditions and does not have ADH activity. Lythraceae (loosestrife) Herbs with 4-6 crumpled petals on the edge of a floral cup (crape-myrtle is a cultivated shrub in this family). Physiological adaptation: Lythrum salicaria has no ADH activity. Nymphaceae (water lily) Herbs with floating round or heart-shaped leaves and large, showy, many-parted flowers. All are in wetland habitats. Morphological adaptation: floating leaves. Oleaceae (olive) Trees or shrubs with opposite leaves and four parted flowers having two to four stamens. Morphological adaptations: buttressed trunks and adventitious roots. Physiological adaptation of Fraxinus (ash): oxidizes the rhizosphere. Poaceae (grass) Herbs with round stems and reduced flowers enclosed in two bracts. Physiological adaptation: Phragmites australis accumulates malate. Salicaceae (willow) Trees or shrubs that are separate sexes with flowers produced in catkins. Morphological adaptations: Populus deltoides has adventitious roots and species of Salix have hypertrophied lenicels, adventitious roots, and spaces in the stem tissues that allow oxygen to reach the roots. Salix nigra Wetlands Research Program Technical Report Y87-1 (on-line edition) Corps of Engineers Wetlands Delineation Manual by Environmental Laboratory http://wetland-delineation.rutgers.edu/87-wetlanddelineation-manual/ Useful supplementary information for making wetland determinations can also be found at the following sites on the World Wide Web: List of wetland plants http://rsgisias.crrel.usace.army.mil/NWPL/ Analyses of normal precipitation ranges and growing season limits http://www.wcc.nrcs.usda.gov/climate/wetlands.html National Wetlands Inventory maps and databases http://www.fws.gov/wetlands/ Table C1 Partial List of Species with Known Morphological Adaptations for Occurrence in Wetlands1 Species Common Name Adaptation Acer negundo Box elder Adventitious roots Acer rubrum Red maple Hypertrophied lenticels Acer saccharinum Silver maple Hypertrophied lenticels; adventitious roots (juvenile plants) Alisma spp. Water plantain Polymorphic leaves Table C2 Species Exhibiting Physiological Adaptations for Occurrence in Wetlands Carex arenaria Carex flacca Carex lasiocarpa Malate accumulation Absence of ADH activity Malate accumulation