Plant stresses and responses De Block et al., Plant J. 41:95 (2005) Plant stresses and responses Plant Physiol Biotechnol 3470 March 21, 2006 Chp 21

Plants are sessile and must deal with stresses in place Plants cannot avoid stress after germination How plants deal with stress has implications in Ecology: Stress responses help explain geographic distribution of species Crop science: Stress affects productivity Physiology and biochemistry: Stress affects the metabolism of plants and results in changes in gene expression www.grainscanada.gc.ca From engineering, stresses cause strains (responses of stressed objects) = changes in gene expression and metabolism in plants Biological stress difficult to define/quantify: What is “normal” metabolism? Limitations to yield? Where on gradient of availability of limiting resource does stress begin? Varies by species, ecotype Heat-stressed wheat

Stresses are abiotic or biotic Fig. 21.1 ABIOTIC STRESSES Environmental, non-biological Temperature (high / low) Water (high / low) Salt Radiation Chemical BIOTIC STRESSES Caused by living organisms Fungi Bacteria Insects Herbivores Other plants/competition Stresses cause responses in metabolism and development Injuries occur in susceptible plants, can lead to impeding flowering, death Ephemeral plants avoid stress Mexican poppies in US desert SW Only bloom after wet winter Die before summer returns Preferable! http://www.geo.arizona.edu/gallery/US/tuc_2.html

Plants must be stress resistant to survive Avoidance also possible by morphological adaptations Deep tap roots in alfalfa allow growth in arid conditions Desert CAM plants store H2O in fleshy photosynthetic stems Stress resistant plants can tolerate a particular stress Resurrection plants (ferns) can tolerate dessication of protoplasm to <7% H2O  can rehydrate dried leaves Plants may become stress tolerant through Alfalfa plant Adaptation: heritable modifications to increase fitness CAM plants’ morphological and physiological adaptations to low H2O environment Acclimation: nonheritable physiological and biochemical gene expression Cold hardening induced by gradual exposure to chilling temps, a/k/a cold-hardy plants Alfalfa taproot www.agry.purdue.edu; www.omafra.gov.on.ca; Heat stressed rose leaf

Let’s walk through one each of important abiotic and biotic stresses View how they affect metabolism Determine how the plant responds to counter the stress ABIOTIC STRESS: Temperature Plants exhibit a wide range of Topt (optimum temperature) for growth We know this is because their enzymes have evolved for optimum activity at a particular T Properly acclimated plants can survive overwintering at extremely low Ts Environmental conditions frequently oscillate outside ideal T range Deserts and high altitudes: hot days, cold nights Three types of temperature stress affect plant growth Chilling, freezing, heat Growth rate Topt Growth temperature

Suboptimal growth Ts result in suboptimal plant development Chilling injury Common in plants native to warm habitats Peas, beans, maize, Solanaceae Affects seedling growth and reproduction multiple metabolic pathways and physiological processes Cytoplasmic streaming Reduced respiration, photosynthesis, protein synthesis Patterns of protein expression Transition temperature Membrane fluidity affects permeability! http://cropsoil.psu.edu/Courses/AGRO518/CHILLING.htm Initial metabolic change precipitating metabolic shifts thought to be alteration of physical state of cellular membranes Temperature changes lipid and thus membrane properties Chilling sensitive plants have more saturated FAs in membranes: these congeal at low temperature (like butter!) Liquid crystalline  gel transition occurs abruptly at transition temperature

Biotic stresses are mitigated by plants’ elaborate defense strategies Buchanan et al., “Biochemistry & molecular biology of plants,” 2001 BIOTIC STRESS: Pathogen (e.g., fungus) invasion Plant reaction to invading pathogens center around the hypersensitive reaction The hypersensitive reaction initiates many changes in plant physiology and biochemistry Defenseless Wild type Early activation of defense related genes to synthesize pathogenesis related (PR) proteins Protease inhibitors to stop cell wall lysis by specific enzymes expressed by pathogen Bacterial cell wall lytic enzymes (chitinase, glucanase) Change cell wall composition Express enzymes providing structual support to cell walls via synthesis of lignin, suberin, callose, glycoproteins Synthesize secondary metabolites to isolate and limit the pathogen spread These include isoflavonoids, phytoalexins Apoptosis at invasion site to physically cut off rest of plant Sequential or parallel events??

How does the plant recognize and defend itself against pathogens? Plant disease has an underlying genetic basis Pathogens may be more or less potentially infectious to a host virulent on susceptible hosts avirulent on non-susceptible hosts Pathogens carry avirulence (avr) genes and hosts carry resistance (R) genes The normal presence of both prevents pathogens from attacking the plant Infection occurs when pathogen lacks avr genes or plant is homozygous recessive for resistance genes (rr) In these cases, the plant cannot initiate the hypersensitive reaction This is bad news! The plant requires this response to survive! Note the communication between pathogen and plant Pathogen: avr genes may code for proteins that produce elicitors bits of pathogen: polysaccharides, chitin, or bits of damaged plant: cell wall polysaccharides Plant: R genes may be elicitor receptors

The hypersensitive reaction initiates a plant immune response The long term plant resistance to a pathogen is similar to a mammalian immune response This is known as systemic acquired resistance (SAR) Secondary metabolites induced by the hypersensitive reaction initiate changes in metabolism in other plant organs through control of signal transduction chains Hours to days: capacity to resist pathogens spreads throughout plant Immune capacity = SAR SAR signaling involves salicylic acid (SA), a natural secondary metabolite SA both induces pathogenesis related gene expression and enhances resistance to infection by plant viruses Fig 21.17

Salicylic acid induces systemic acquired resistance Fig 21.18 volatilized Local SA production induces distal production and SAR via SA transport in xylem Methylation into MSA, volatilization and distal detection High constitutive SA levels result in plants with high ability to withstand pathogens Mechanism by which SA induces SAR unknown Jasmonic acid also mediates disease and insect resistance JA also mediates other developmental responses: PGR? All stress affects photosynthesis: productivity and survival Knowledge of how stress is perceived and transduced central to understanding plant metabolism