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Transcript
Desiccation Tolerance Mechanisms and Evolution Mel Oliver and Brent Mishler Desiccation-tolerance. The ability to revive from the air-dried state (the air being of low relative humidity) thus experiencing protoplasmic dehydration without suffering permanent injury Bewley and Krochko. 1982 Types of Desiccation-tolerance. Plants whose tolerance to water loss is low. Plant structures that are adapted to withstand desiccation and for which water loss is an expected event. - seeds. Plants that are capable of tolerating desiccation regardless of the rate at which water loss occurs. Plants that are capable of tolerating desiccation only if water loss is a slow process. Desiccation-tolerant Plants. Desiccation-tolerant ALGAE LICHENS BRYOPHYTES Modified Desiccation-tolerant FERNS ANGIOSPERMS Distribution of Desiccation Tolerance in the Plant Kingdom ferns Equisetum Selaginella cycads conifers gnetophytes Angiosperms Gingko Isoetes Lycopodium mosses hornworts liverworts Seed Plants Tracheophytes Land Plants Oliver, Tuba and Mishler 2000 Tortula ruralis Selaginella Selaginella bigelovii Polypodium virginianum Orthodox Seeds Photo Courtesy of Dr Christina Walters USDA NSSL Fort Collins Distribution of Desiccation tolerance in the Angiosperms Cyperaceae Labiatae Poaceae Gesneriaceae Velloziaceae Liliaceae Magnoliales Hamameliales renunculids Oliver, Tuba and Mishler 2000 Angiosperms Scrophulariaceae Xerophyta villosa Myrothamnus flabellifolia Photo Courtesy of Dr. Jill Farrant and Clare Vander Willegen University of Cape Town SA Craterostigma wilmsii Xerophyta viscosa Sporobolus stapfianus Craterostigma plantagineum Hydrated Dry Rehydrated Photos Courtesy of Dr. Dorothea Bartels University of Bonn Critical Parameters for Desiccation-tolerance. Limit damage to a repairable level Maintain physiological integrity in the dry state Mobilize repair mechanisms upon rehydration Bewley 1979 Essence of Desiccation-tolerance. Testable Hypothesis Cellular Repair Cellular Protection Bryophyte Model RAPID WATER LOSS Constitutive Cellular Hormone ? Protection Induction of Recovery and Repair Mechanisms Hydrated Dry Rehydrated Angiosperm Model SLOW WATER LOSS Induction of ABA Cellular Protection Re-establishment Processes Hydrated Dry Rehydrated Postulated Evolutionary History of Desiccation Tolerance in Land Plants Equisetum Selaginella ferns cycads Angiosperms conifers gnetophytes Inducible protection (repair?) and later poikilochlorophylly Gingko Isoetes Lycopodium mosses hornworts liverworts S Developmentally programmed protection - propagules T Inducible protection plus repair? Developmentally programmed protection - spores? Constitutive protection and repair Oliver, Tuba and Mishler 2000 Loss of vegetative desiccation tolerance in the ancestral lineage Bryophyte Model RAPID WATER LOSS Constitutive Cellular Hormone ? Protection Induction of Recovery and Repair Mechanisms Hydrated Dry Rehydrated Tr 288 Phylogenetic Gene Search Dhy Tr 288 Gene In 1 A B C D E F G H I J K L MN O In 2 Expected PCR Products (Sequence for GPN-Box Consensus primers Identity) E F G H Unrooted Tortula Phylogenetic Network Probable tree root Occurance of Tr288 Orthologs Calyptopogon 288 Tortula papillosa 288 Tortula sinensis 288 288 Tortula muralis Tortula andersonii 288 Tortula indet NSW 288 288 Tortula handelii Tortula amphidiacea 288 Tortula subaristata 288 288 Tortula ruralis Tortula caninervis 288 Tortula cavelii 288 Unrooted “Deep Green” Phylogenetic Network Occurance of Tr288 Orthologs Riccia frostii Riccia membranacea Riccia albolimbata Riccia sulivantii Riccia atromarginata Targionia Riccia albida Asterella Blasia Haplomitrium Notothylas Megaceros Lunularia algal ancestor Equisetum Sphagnum palustre Exostratum Octoblepharum Huperzia 288 Isoetes Polytrichum piliferum Buxbaumia Polytrichum commune Funaria Grimmia 288 Leucophanes Calyptopogon 288 Arthrocormus Tortula ruralis 288 288 Mitthyridium Tortula princeps Tortula muralis 288 288 288 288 Sequoia Selaginella Anthoceros fusiformis Tetraphis Pterogonium Osmunda Angiopteris Psilotum Anthoceros Sphagnum cuspidatum Probable Network Root Lophocolea 288 288 Phylogenetic Approach to Functionality Establishment of a correlation between the presence of a gene and a specific phenotype Establishment of the role of a gene in the evolution of a particular phenotype Establishment of the importance of a particular mechanism in the evolution of a particular phenotype, e.g., induced repair upon rehydration versus induced protection during drying in desiccation-tolerance Collaboration with Brent Mishler - UC Berkeley Deep change in function A phylogenetically distant comparison = large background differences Recent Change in function A phylogenetically close comparison = low background differences Increasing complexity Ancestor-descendant comparison using reconstructed ancestral states