MELBOURNE BIOLOGY NETWORK MEETING 28.5.02 SPEAKER: Dr Don Gaff, Monash University (retired). Survival in Extreme Conditions – Resurrection Plants Resurrection plants: * leaves, shoots and roots survive air-drying. Water content drops to 4 – 8% dry weight. * drying takes 3 – 7 days after rain. * rehydration takes 12 – 24 hours * air-dried leaves survive 2 – 5 years * there is no limit on the number of hydration/rehydration cycles that RPs can survive. Abscisic acid (ABA) * in plants generally [ABA] with stress. There is a 40 – 100 times increase in situations such as: drought, salt, cold, heat, low levels O2. * ABA treatment increases drought resistance of these plants in general: root growth closes stomata shoot growth leaf hairiness leaf senescence (reduces number of leaves). o Loss of chlorophyll o Leaves shed (possibly through ethylene action). Is ABA involved in Resurrection Plants? If a small stem of Borya (RP) is air dried, it can be resurrected. If a leaf is cut from the plant and dried quickly then it dies but the chlorophyll stays. If such a detached leaf is dried slowly over two days, chlorophyll is lost and the plant stays alive. [ABA] increases in Borya leaves as they dry. Also ABA treatment of wellhydrated Borya leaves allows them to survive subsequent drying. Cf. Leaves of the South African resurrection grass Sporobolus stapfianus can survive so long as they stay on the parent plant. If detached leaves are dried they will always die. The basic hypothesis: there must be a signal from the root system to the shoot. (8) Physiological changes Response (7) Specific protein mRNA (6) Activation of specific gene (5) ABA-induced compounds Signal transduction (across cell to nucleus) (4) ABA receptor on cell membrane Leaf Root (3) ABA rises in xylem (2) ABA secreted into root xylem Transport of hormone Perception stimulus (organ response) (1) Drying soil Please note that some hormones (eg. ABA & gibberellins) move through xylem as well as phloem. It is the [ABA] at the cell membrane (cell wall space) that is important rather than the [ABA] in the cells in inducing stomatal closure. Some evidence indicates that detached leaves can produce their own ABA (eg. Borya and Sporobolus, see above). Then researchers started looking for another substance as a root signal. No singal plant hormone gave enough improvement in drought tolerance of Sporobolus leaves. Perhaps a particular mixture of several hormones is required, cf. Experiments in South Australia with grape vines. An experiment was done using split-root irrigation of a grape vine. The root of a plant was split and watered on one side only using drip irrigation with the other side remaining dry. The following was found: The water content in the foliage was the same. The stomata closed. 2 There was less growth of foliage. Grape yield was reduced only a little. Grape quality was much improved. This is good news for reducing irrigation volumes. No single hormone explains the postulated root signal. As S. stapfianus (RP) is dehydrating the number of proteins produced in the plant is increased. No explanation yet. In a non-resurrection plant often studied by geneticists: Arabidopsis thaliana – the Japanese showed that there are four different drought-induced pathways that stimulate suites of genes (two pathways with ABA). The pathways involve “transcription factors” that activate the genes. Hormones such as ABA, auxin, ethylene and gibberellin stimulate the transduction pathways (which can interact) and the system amplifies the response. Waterlogging (Plants generally) In waterlogged soil plants: reduce [gibberellin] and [cytokinin] in the roots as well as increase [ABA] and [IAA] in the roots. This leads to an increase in the root/shoot ratio of cytokinin and IAA. Thus root growth is reduced. Low cytokinin / high IAA tends to stimulate root growth. Production of ethylene: Methionine SAM Step stimulated by low O2 ACC synthetase ACC (transported through plant) ACC oxidase Step impaired by low O2 Ethylene (C2H4) SAM = S-adenylmethionine ACC = 1-aminocyclopropane-1-carboxylate Ethylene acts where it is produced so, by definition, it is not a hormone. That is, it is not produced in one area and transported to another. Nevertheless released as a gas it can cause growth response in other organs in other 3 plants. Moreover, in combination with precursor ACC (transportable) it forms part of a hormone-type system. ACC is formed in waterlogged roots and transported up in the xylem. When the ACC reaches aerated tissues, the oxygen present allows conversion of ACC to ethylene. Growth of new roots is stimulated in the aerated by the ethylene in combination with a favourable cytokinin/IAA ratio. In the new roots (in the aerated soil) most of the root cortex collapses and large gaps are produced that fill with gas that allows a better supply of air. Ethylene (from the ACC from the lower waterlogged roots) causes this breakdown of the cortical cells. This is tissue with the large gaps is termed aerenchyma. Some ‘recent’ references on plant hormones 1st Year Uni: Knox B, Ladiges P, Evans B, Saint R (2002) “Biology” 2nd ed. McGraw-Hill, Roseville NSW Ch 24 pp 630-647 2nd/3rd Year Uni: Salisbury F, Ross C (1992) “Plant Physiology” 4th ed. Wadsworth, Belmont, Calif. Ch 17-18 pp 357-406. Atwell B, Kriedemann P, Turnbull C, eds. (1999) “Plants in Action” Macmillan, S Yarra. Ch 9 pp 284-306. Starr C and Taggart R (2001) “Plant Structure and Function” 9th ed. Thomson/Brook/Cole. Anderson J and Beardall J (1991) “Molecular Activities of Plant Cells”. Blackwell, Oxford. www.plant-hormones.bbsrc.ac.uk/education/kenh.htm www.biologie.uni-hamburg.de/b-online/e31/31h.htm 4