Water, drought, adaptation

From presentations by Marek Neuberg and Jan Pokorn compiled by Jan Kvt

Daily course of transpiration

Effect of stomatal regulation on the daily course of transpiration in plants growing with different water supply, from 1 to 5 (www.ictinternational.com.au)

Control of transpiration

Pevzato z: http://www.mhhe.com/biosci/pae/botany/vrl/images1.htm (20.6. 2004)

Water flows within the plant along a water potential gradient

Water flow in plant cells is a passive process. That is, water moves in response to physical forces, towards regions of low water potential. Consequently, there are no metabolic pumps for water movement (such as reactions driven by ATP) that would push water from one place to another.

Aquaporins increasing cell membrane permeability

There are proteins in the cell membranes of many plant cells, and these proteins are sensitive to the water status of the cells: they increase the overall permeability of water flow through the cell membranes

What happens when the external solute concentration is increased?

Increasing the solution concentration to 0.3 M sucrose causes the cell to lose water, to the point where the membrane is just about to pull away from the cell wall (called incipient plasmolysis) The increased solute potential of the external solution draws water out the cell, reducing the cells turgor pressure Again, water moves from a high to a low water potential.

Water potential in plants

Water potential strongly affects the growth of cells and photosynthesis in plants. Thus, like measurements of body temperature of humans, water potential is a good overall indicator of plant health. Plant physiologists know a wide variety of ways to measure water potential in plants (e.g., plasmolysis, pressurizing of leaves)

Plant water status and the operation of physiological processes

Because of transpirational water loss, plants are seldom fully hydrated Plants thus readily cope with conditions of negative pressure, or water deficit, up to a point where different physiological processes become inhibited Thus by measuring the water potential of plants we can get an idea of how stressed they are.

The whole plant water transport network

Transport of water through the whole plant involves several interacting components, each with different driving forces and rates of transport We will consider the following components of water transport from roots to shoots: Soil transport properties (remember plants are in contact with the soil!) Root hydraulic transport, water flow across roots Water flow within the xylem (stems and leaf veins) Evaporation (transpiration) from the leaf intercellular air spaces to the atmosphere

Soil water transport: an overlooked dimension

Water supply to the plant is critically
influenced by how water moves through the soil; what are the driving forces for water flow in soil and which soil characteristics affect water flow?

Soil type and texture determine how easily water flows in the soil by affecting the soil porosity:
Sandy soils have large particles and large pores in between (1 mm or more) Water IS NOT held tightly by this soil Clay soils have much smaller particles and pores in between (~2 microns) Water IS tightly held by this soil

How does water move in soil?

Water moves through the soil predominantly by bulk flow, that is down a pressure gradient under NO influence of solutes As plants absorb water from the soil, they deplete the soil of water near the surface of the roots. This reduces P of the water near the root surface and establishes a pressure gradient from the soil into the root. Because water-filled pore spaces of the soil are INTERCONNECTED, water moves through the soil by bulk flow down a pressure gradient.

Water uptake by roots; the gatekeepers of whole plant water flow

Water enters the plant typically first at the root level Water moves in the root through several interconnected and complicated pathways Unlike the soil where water moves only by bulk flow, water flow through the root is more complicated and occurs by osmosis Water flow through roots takes two major pathways through the epidermis and cortex of the root towards the central cylinder with vascular bundles: Apoplastic Symplastic The relative importance of one or the other path is not clear.

Pathways of water movement in roots

Apoplastic and symplastic paths

Root pressure and guttation

Root pressure occurs when soil water potentials are very high and transpiration rates are low If transpiration rate is high, water is taken up so rapidly that a positive pressure cannot develop Plants that produce high root pressure frequently form liquid droplets on the edges of their leaves; a process called guttation Positive (root) pressure causes the water exudation

Water transport in the xylem

In most plants, the xylem constitutes the longest part of the whole plant water transport pathway For a 1 m tall plant, more than 99.5% of the water transport pathway will occur within the xylem. Compared to the root, xylem water transport is a fairly simple low resistance transport pathway. We will consider now: The role of negative pressures in xylem transport And the importance of safe xylem transport under negative pressure

Negative or positive pressure driven water flow?

Pressure gradients needed to move water through the xylem could come about in two ways: Positive pressures: - Hypothetically positive pressure could drive water to the tops of trees, but - Root pressures are generally less than 0.1 MPa (lift water only 1 m above the ground) and pressures of at least 2 MPa are required to supply some of the tallest trees on the planet - Root pressure is energetically expensive... - Also, evaporation will easily collapse positive pressure gradients... - Negative pressures: - Solar power! Plants use the evaporative power of solar energy to pull water through the vascular system. - So, water at the tops of trees develops a large tension to pull water upwards through the xylem

Xylem water transport under tension is risky

The large tensions that develop in the xylem of trees can create some problems: First, tension results in an INWARD pull on the tracheary cell system. The development of lignified walls is necessary to allow resistance to implosion from this force... The next problem is that water is METASTABLE and very sensitive to slight changes in gas content Recall that pure degassed water is very strong, but with gas added, the water column can become increasingly easily broken

Metastable state of water under tension

The other problem is that water is METASTABLE and very sensitive to slight changes in gas content Recall that pure degassed water is very strong, but with gas added, the water column can become increasingly easily broken As tensions increase, there is an increased tendency for air to be pulled in from microscopic pores in the xylem wall that contain air (from respiring living cells or just close to lenticels, etc.) This is called air-seeding.

Drought survival

Adaptation
Plants optimize their dispersal, probably by root competition so as to obtain the maximum amount of growth per unit rainfall.

Water stress due to water shortage:

Mostly combined with stress due to excessive irradiance and overheating, often also with stress due to high soil salinity

Possible reasons of water shortage:

High rate of evapotranspiration at low rainfall

Water bound osmotically in soil (saline soils) Frozen soil

Thin soil layer, underdeveloped root system

Symptoms of water stress in plants?

Reduction of cell volume, dehydration of protoplasm Decreased turgor and slower growth, esp. elongation Metabolic changes: osmotically active substances, etc. Disturbed protein metabolism and aminoacid synthesis Slower meristematic cell divisions, anomalous meiosis Enhanced synthesis of abscissic acid closure of stomata Changed allocation of assimilates, increased root/shoot ratio Characteristic morphogenetic alterations

Osmoregulation and stress metabolites

Cumulation of osmotically active substances Maintenance of turgor Non-structural saccharids (carbohydrates) and aminoacids Stress metabolites accumulate in cytoplasm as a result of stress (betaines and the aminoacid proline) Proline Its source can be either the synthesis from glutamate or hydrolysis of proteins Heat shock proteins

Water stress
Poikilohydric plants, which maintain an equilibrium between atmospheric humidity and the hydrature of their protoplasm (algae, fungi, lichens and bryophytes) Homoiohydric plants - hydratation of protoplasm is independent of air humidity (higher plants) Xerophytes (irrespective whether poikilohydric or homoiohydirc) which are non-succulent and nonhalophytic, obtain their water supply from local precipitation and atmospheric moisture. Therophytes vs. Geophytes - two adaptations to drought Time of flowering and fruiting.

Poikilohydric xerophytes
Plants of this type lose water rapidly in the dry season and survive the adverse period in a state of intense desiccation. Parkinsonia microphyla 250 years of drought

Grimmia pulvinata drkavka podukovit

Bazzania stolonifera -

Jtrovka rohozec
Tortula ruralis rourkatec obecn

Ceterach officinarum kyvor lkask

Selaginella lepidophylla

The greatest numbers of poikilohydric angiosperms occur in South Africa: various species of Scrophulariaceae, Myrothamnaceae, Velloziaceae, Cyperaceae, Poaceae

Myrothamnus flabellifolia

Vascular plants both dehydration and rehydration must be slower After 2-3 years of almost absolute dryness they can completely recover, damaged root tips and trichomes quickly regenerate Ramonda myconi Ramonda serbica

Homoiohydric xerophytes 1
Malacophyllous species (soft-leaved) which are characteristic of semi-desert conditions where winter or seasonal rains ensure periodic alleviation of drought. When exposed of drought, they shed their leaves. Cistus, Lavendula, Rosmarinus

Homoiohydric xerophytes
Malacophytes Can live under semidesert conditions Their leaves wilt, or are shed off during
the dry period

Cistus, Lavendula, Thymus, Rosmarinus, Artemisia, etc.

Rosmarinus officinalis Lavendula officinalis

Homoiohydric xerophytes 2
Sclerophyllus species (hard-leaved) which are able to maintain a favourable water balance by reducing their transpiring surface area during predictable drought periods. Well-developed root system. Quercus ilex, Pinus pinea, Spartium junceum

Sclerophytes

Plants with tough leaves or entirely leafless Reduced transpiring surface Leaf turnover of several years Extensive and deep root systems

Araucaria araucana leaves persist up to 25 years

Quercus ilex

Pinus pinea Schinus molle

Spartium junceum Vtenk stinolist

Olea europaea

Succulent xerophytes
Water-storing species Crassulacean acid metabolism (CAM) Nananthus, Conophytum, Opuntia

Succulent xerophytes
Water accumulation during (mostly rare) wet periods. Shallow-rooted, sometimes lacking roots, e.g., epiphytic tillandsias.

Opuntia puberula

Or their roots die off during dry periods and regenerate when water is again available.

Succulents CAM metabolism (Bromeliaceae, Orchideaceae, Cactaceae, Aizoaceae, Euphorbiaceae) Leaves cylindrical, sphaerical, often plants leafless Photosynthesis rather in stems Reduction of number of stomata, thick epidermis, stomata immerged

Desert conditions: long periods without water, some plants completely buried in soil
Efficient control of water output (loss) Lithops francisci

Lithops salicola

Drought tolerance 1
The maintenance of sufficient physiological integrity so that metabolism can be reactivated quickly upon rehydration. A repair mechanism that can be put into effect on rehydration and which can repair any damage caused to membranes and membrane-bound organelles.

Drought tolerance 2
Density of protoplasm (sugars), waterstress resistant proteins? Seeds have waterrepellent layer(s); importance of hardness and morphology of seed coat

Reduction of water loss 1

In a drying out soil profile, roots not only grow faster than in a well-watered one but also achieve a substantially greater total length. Aerial roots of epiphytic orchids.

Specially adapted water-absorbing leaf

hairs (trichomes) of the Bromeliaceae Stomatal pores

Tillandsia usneoides ( Spanish moss)

Reduction of water loss 2

Cuticular adaptations to water stress Osmoregulation and stress metabolism, proline accumulation

Reduction of water loss 3

Diurnal variation in CO2 uptake, PEP carboxylase, CAM metabolism

Conclusion
The principle of optimization with regard to water loss can be summarized as balancing the risks of water output against the benefits of carbon fixation.

Thank you and see you again!