Themes > Science > Botanical Sciences > Plant Hormones, Nutrition, and Transport > Nutrition, Transport, and Bioregulation in Plants

Introduction

Photosynthesis

Photosynthesis is the conversion of solar energy by plants into several forms of chemical energy. (2) Photosynthetic productivity is the carbon balance of a plant over a time period and depends on the following external environmental factors: light availability, water (including air humidity), nutrient availability, carbon dioxide availability, and temperature.

Internal Factors Affecting Plants

Internal environmental factors include leaf age and chlorophyll content, osmotic adjustment, the presence of strong sinks, photorespiration, and plant hormones

External Factors Affecting Plants

Photosynthetic productivity is the carbon balance of a plant over time and depends on the following factors:

  • light
  • water
  • nutrients
  • gases
  • temperature

Radiation

Types of Radiation and Their Effects

  • Electromagnetic Spectrum (16-2)
  • UV can alter molecular structure
  • Photosynthetically active Range (360-760 nm)
  • Infrared has little energy (accelerates molecular reactions)

Energy From Radiation

Radiation energy is described by the following equation:

E = hc/l = hn

  • E = energy of a photon
  • h = Planck’s constant (= 6.63 x 10-34 J s)
  • c = speed of light (= 3 x 108 m s-1)
  •    = wave length (=cm)
  • v   = frequency of oscillation

Photosynthetic Efficiency

  • 0.35 g C m-2 d-1 in average ecosystems
  • average efficiency is 0.14%
  • theoretical is 3.2%

Factors Affecting Photosynthetic Efficiency

Interception of light is determined by Beer’s Law

I = Ioe-kL

Io = irradiance at the top of leaf canopy

I = irradiance at a point in the canopy above which there is a leaf area index (L)

k = extinction coefficient (empirical)

leaf sizes and shapes

leaf orientation

light intensity

leaf reflection and absorption (16-3)

Phytochromes

Light responses are caused by phytochromes which exists in two forms Phytochrome interconversions (16-6) Phytochrome Prosthetic Group (16-7)

Inductive - photoreversable (e.g. leaf closing)

High Irradiance Responses (HIR) (longer response, e.g. stem elongation)

Phototropism (16-8)

Phototropism refers to the directional alterations in growth that occur in response to directional light stimuli, for example, the orientation of leaves in relation to the sun. One mechanism for controlling these alterations is indoleacetic acid.

Photonasty

Photonasty refers to the reversible light movements and related phenomena that occur in response to directional and non-directional light stimuli, ie. the opening and closing of flowers. It is caused by turger changes at the base of leaflets.

Photoperiodism

Photoperiodism is the non-directional developmental responses to non-directional but periodic light stimuli.

Signal for flowering (16-9)

  • red or white flash inhibits flowering of short-day plants and induces flowering of long-day plants
  • far-red lignt flash reverses effect of exposure to red flash

Photomorphogenesis

Other non-directional developmental responses to non-directional and non- periodic light stimuli

Seedlings grown in darkness are elongated and pale (encourages plant to elongate to find light).

Stem and leaf expansion are light sensitive.

Fluorescent and incandescent light of equal intensities causes differences  (16-10) in growth .

Sensitivity to light quality is important factor in adaptation to shade (red/far-red is a signal for plant density.

Water

General Comments About Water And Plants

Plants are typically about 90% water.

A typical crop or grassland will transpire about 500 kg of water per kg dry wt. produced.

Water conveys inorganic nutrients and photosynthetic products to various parts of the plant.

Water is also the electron donor for photosynthesis.

Water evapotranspiration also keeps plants from overheating.

Factors Influencing Movement Of Water In Plants

  • Transpiration
  • Water potential
  • Movement of water in capillaries
  • Coupling transpiration with absorption

Transpiration

Stomatal pores (16-11) (photo of pores) (16-12) in leaves open to allow movement of carbon dioxide in for photosynthesis

Water vapor is lost through pores by transpiration (mechanism follows Fick’s Law of Diffusion)

Loss of water from non pore areas is restricted by a waxy impermeable cuticle

Control by Stomata

When water stressed, plants increase thickness of cuticle

In light, guard cells accumulate potassium ions and organic acids

This decreases their osmotic pressure which causes them to fill with water and enlarge the stomatal pores

Extra water losses will cause the cells and pores to shrink, reducing water loss

Under conditions of water stress, leaves produce a hormone, abscisic acid, which promotes stomata closure

Stomatal pore size can also be regulated by CO2 (16-13) concentration

Absorption

Absorption refers to uptake of water by roots to compensate for water losses by transpiration

During daylight, transpiration exceeds absorption and cells shrink lowering their water potential

At night, stomatal pores close and water potential of leaves (16-14) becomes restored.

Movement of Water in Capillaries

A typical xylem vessel has a radius of 20 um which has a capillary rise of 0.7 m

Pores in the polysaccharide matrix of cell walls have radii of 5 nm which are able to support a water column of 3 km

Some estimate that the tensile strength will support continuity of a water column of more than one mile

Water Potential in Plants

Water potential (16-15) was discussed in the previous lecture

The major components in plants are turgor pressure and osmotic potential

Turgor pressure is the pressure difference inside and outside cell usually positive

  • 0.5 to -3.0 MPa in transpiring leaves

Osmotic Potential

Describes effect of solutes on diffusion properties of water (16-16)

In soils, is influenced by surfaces and capillary spaces

Dissolved salts in soil have minimal influence on OP in plants

Wilting point = lower limit of water availability for plant (about -1.6 MPa)

Matric Potential

Matric potential refers to the effect of porous solids on water movement

Like plant tissue, soils have pores and charged surfaces which oppose movement of water into plants

These become less significant when soil is wet or saturated

Summary of Factors Influencing water potential

  • Hofler-Thoday Diagram (16-17) shows the change in water potential and its components as a cell loses water

Effect of Drought on Plants

Water loss exceeds absorption and plants become dehydrated

Photosynthesis is reduced when water potential is -1 to -3 MPa

Stomatal conductance is limited reducing water loss and CO2 entry related to hormone abscisic acid (ABA)

Effect on thylakoid bound reactions: low water reduces fluorescence, electron transport, and photophosphorylation

Dark Reactions (CO2) fixation is limited

Photoinhibition occurs because photosynthesis is limited

Reduced plant growth

Modification in development and morphology

Reproductive development

Drought Tolerance Mechanisms: Avoidance of Plant Water Deficits

Drought escape - short growth cycle, dormant periods

Water conservation - small leaves, limited leaf area, stomatal closure, high cuticular resistance, limited radiation absorption

Protective solutes (sugars, alcohols protect cytoplasmic proteins), desiccation tolerant enzymes

Turgor maintenance - osmotic adaptation (increase in solutes), low or high elastic modulus

Efficient use of available water (stomatal closure, leaf rolling)

Maximal harvest index (optimizing water use and yield of desired portion of plant)

Responses to Nutrients

General

Forms of Nutrients Used by Plants

N: NO3-, NH4+, or N2

P: H2PO4- or HPO4=

S: SO4=

Others: K+, Ca++, Mg++, Fe++, Fe+++

Absorption of Nutrients

Absorption nutrients is independent of water absorption

Active transport is usually involved as the concentration in the plant is usually higher than outside

Once inside, nutrient concentrations are reduced by use or transport to other parts of plant

Absorption follows Michaelis-Menten kinetics

Role of Microorganisms in Nutrient Uptake

Release nutrients via mineralization

Dissolve nutrients from insoluble ores, release adsorbed forms by lowering pH

Direct influence on nutrient uptake (mechanism unknown)

Competition with plants for nutrients

Nitrogen fixation

Transport of Nutrients

Inorganic nutrients are transported from the roots to leaves primarily via xylem

Photosynthate is transported to the various plant parts via phloem

Pressure flow hypothesis for explanation of rapid mass transfer of organic solutes in phloem (16-23)

Responses to Gases

Plants Interact Primarily with Three Gases:

CO2 - carbon source

O2 - product of photosynthesis

N2 - nitrogen source

Movement of Gases

Gases move in and out of guard cells

Gases move freely once inside tissue

Large tissue surface area enhances gas movement

Lenticel: Loose patches of cells through bark facilitate gas transfer in stems

Roots freely exchange gases

Flooded crops send out aerial roots

General Responses to Carbon Dioxide

Atmospheric CO2 has increased from 280 to 350 umol mol-1 in past century caused by fossil fuel combustion

threat of global warming (16-24)

Increasing CO2 (double) increases photosynthesis (16-25) 35-50% in C3 plants but not C4 plants. Increase is short term and not sustainable

Responses to Elevated Carbon Dioxide Levels

Stimulates stomatal pore closure and aperture and decreases stomatal densities

Increases the ratio of photosynthesis to transpiration, thus increasing water use efficiency

Increases temperature effects, nutrient use efficiency, and light effects at high intensities

Responses to Gaseous Dinitrogen

Nitrogen-fixing bacteria (Rhizobium) (16-26) in nodules of legumes fix nitrogen and grown on organic metabolites synthesized by plant host

Nitrogen fixation is anaerobic

Responses to Temperature

General

Survival Range is -89 to 70oC

Growth range is >0 to ~40oC

Physiological basis for control

Transpiration keeps plants from overheating

Plant tissue is a poor conductor of heat (16-27)

Temp. optima of different plants (16-28)

Effects on Plant Development

Extension of thermal time - seed germination time is decreased by increased temperatures

Degree-days - accumulative number of days above a certain base temperature

Vernalization plants - require seasonal periods of temperature highs and lows for proper growth and reproduction may be quantitative or obligate (e.g. winter wheat)

Dormancy and leaf abscission - plants may have winter or summer periods of dormancy (also affected by light)

Effects of High Temperatures

Synthesis of high shock proteins (short term protection)

Protein denaturation

Loss of membrane integrity

Ion leakage

Plants can adapt to temperature extremes; mechanism unknown

Effects of Low Temperatures

Chilling injury in tropical or subtropical plants: inhibited growth, germination, and reproduction

Damage to cell membranes and electrolyte loss

Ice crystals form in tissue causing membrane damage, electrolyte loss

Concentration of cell solutes, solute precipitation, protein denaturation

Mechanisms of Frost Tolerance

Lowering of osmotic potential

Increased levels of carbohydrates (compatible solutes)

Production of abscisic acid

Supercooling: lowering of freezing point of tissue-associated water

Plant Hormones

Hormone (16-29) 

  • chemical secreted in one part of plant that moves to other plant parts and acts on specific target cells

Auxin (16-30)

  • hormone that affects growth of different plant tissues

Gibberellins

  • lipid soluble; can cross membranes
  • break seed and bud dormancy
  • induces amylase production in germinating seed embryos
  • stimulate biennials to flower during the first season
  • stimulate fruit set

Cytokinins

  • differentiation of new cells
  • conversion of proplastids into functional chloroplasts
  • promote fruit development
  • help retard senescence (aging)

Abscisic acid

  • induces dormancy in buds and seeds
  • induces reduced cell division
  • signals changes related to drought

Ethylene

  • stimulates ripening of fruit, called climacteric (caused by rapid CO2 rise and fall)
  • leaf abscission
  • stimulates radial growth of stems and roots
  • breaks dormancy in buds and seed
  • stimulates flower production


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