You are on page 1of 8


is emitted by the Sun and enters the biosphere. Through photosynthesis this radiation energy is fixed in plants in the form of latent chemical energy, from which all links in the food chain derive the energy to carry on vital processes. Thus, radiation is the primary source of energy for the synthesis of organic material, and by regulating the heat and water balance of the earth it provides the necessary energy for life on earth. For plants however, solar radiation is not only a source of energy (Photoenergetic effect), but it functions also as a stimulus for developmental processes (Photocybernetic effect) and occasionally as a stress factor (Photoinhibitory or photodestructive effect). Each of these effects is triggered by the uptake of light quanta and every radiation dependent process is mediated by highly specific photoreceptors, whose absorption spectra correspond to the action spectra of the respective photobiological events (table 1.6). The timing, duration, direction of incidence and spectral composition are important factors of the incoming radiation. The amount of radiant energy reaching different locations on the earths surface depends on their positions with respect to the sun. As a result of the rotation and revolution of the earth, the input of solar energy is periodically changing, and thus imposes specific climate patterns upon all terrestrial phenomena. The periodic alternation of day and night is the astronomical trigger responsible for regulating diurnal and seasonal biological rhythms. Moreover, solar radiation, by acting as a signal, controls many developmental processes such as germination, directional growth and the shaping of external form.


The biosphere receives solar radiation at wavelengths ranging from 290 nm to about 105 nm. Radiation at shorter wavelengths is absorbed in the atmosphere by the stratospheric ozone and oxygen in the air; the

limit of the longer wavelengths is determined by the concentration of water vapor and carbon dioxide in the atmosphere. On average 45% of the incoming solar radiation falls within the spectral range 380-710 nm, this is the range of wavelengths utilized by plants for photosynthesis (photosynthetically active radiation, PhAR; often referred to as the 400-700 nm range). At the lower end of the radiation is ultraviolet radiation (UV-A 315-380 nm and UV-B 290-315 nm) and at the upper end, is infrared radiation (IR 750-4000 nm; see table 1.6). Additionally, plants receive and emit thermo radiation (long wave IR 4000 to 105 nm).

PHOTOENERGETIC EFFECT ON PLANTS Photoenergetic effect is the effect of solar radiation as a source of energy. Biological energy conversions can be categorized into two groups: i) photosynthesis (the process whereby solar energy is fixed to yield energy useful to organisms and industry), and ii) biomass conversion (the product of photosynthesis) into energy. Photosynthesis occurs in plants, algae and photosynthetic bacteria, while biomass conversion reactions often occur in non-photosynthetic microorganisms. Photosynthesis is often regarded as a CO2 anabolic reaction, whereby glucose is formed from CO2 and water. CO2 anabolism is an energyconsuming reaction in that it utilizes chemical energy produced by

photosynthesis. In its narrowest sense, photosynthesis can be regarded as a process whereby energy is supplied for CO2 anabolism. In a broader sense, photosynthesis, including CO2 anabolism, can be divided into several steps: i) photoelectric charge isolation using photon energy (conversion to electrical energy), ii) fixation of electrical energy in the form of chemical energy (ATP synthesis), and iii) chemical reactions involving ATP (fixation of CO2, and hydrogen production).

The supply of energy for CO2 anabolism is common to all photosynthetic organisms which exhibit photosynthesis. Energy conversion, ATP synthesis and the production of both CO2 and hydrogen on the other hand, are not unique to photosynthetic organisms, but occur in all types of microorganisms, and are in fact similar to the respiratory processes which occur in mitochondria of higher organisms. Two types of photosynthesis are distinguishable on the basis of source of the electrons used as energy carriers. In plants such as green algae, and cyanobacteria (blue-green algae), water is the electron source, while in photosynthetic bacteria, organic or sulfur compounds provide electron sources. Photosynthetic mechanisms which occur within plant photosynthetic membranes are schematically presented below. Two Photosystem II water molecules are initially decomposed by four incident photons, to yield one oxygen molecule and four excited electrons. Excited electron energy is subsequently utilized in ATP synthesis. Unlike in the case of ordinary chemical reactions, ATP synthesis cannot be stoichiometrically analyzed. The ratio of excited photons to ATP produced is still a somewhat debatable issue. Although it has generally been thought that two photons give rise to the formation of two ATP molecules, some researchers claim that three photons are involved. Furthermore, other researchers have suggested a loose coupling between proton transport and ATP synthesis: Subsequent to their energy release in ATP production, Photosystem II electrons are transported to photosystem I, where they are again excited to a higher energy level, allowing them to be utilized for NADP reduction. NADP serves both as an electron carrier and an oxidizing and reducing agent in vivo. Two photons are utilized per molecule of NADP reduced.

Experimental data indicates that between 8 and 12 photons are required for fixation of one molecule of CO2. Since the energy equivalent of one photon (700 nm) is approximately 170 kJ/E, and the change in free energy during the fixation of CO2 is approximately 450 U/mol, the energy efficiency of this process for monochromatic light of a wavelength of 700 nm is estimated to be approximately 21-33%.

Schematic representation of mechanisms involved in plant photosynthesis However, owing to the quantum nature of photosynthetic reactions, energy efficiency decreases if light of shorter wavelengths (i.e. higher quantum energy) is used. Additionally, energy losses, energy requirements for plant growth, and the distribution of solar energy wavelengths need to be considered. Plant photosynthesis takes place only in the presence of visible light (400-700 nm). However, solar light contains both visible and infrared components. Since visible light accounts for about 45% of all solar energy, the maximum achievable energy efficiency for CO2 fixation using solar radiation is approximately 13%.

PHOTOCYBERNETIC EFFECT ON PLANTS Photocybernetic effect refers to the functions of solar radiation as a stimulus for developmental processes. Light and heat are the two main components of radiation and both play a vital role as a stimulus for growth in plants. Heliotropism Heliotropism is the diurnal motion of plant parts (flowers or leaves) in response to the direction of the sun. Heliotropism was first described by Leonardo da Vinci. The term "heliotropism," though, was introduced in the early 1800s by A. P. de Candolle, for the growth of the stem tip towards light, which is now called phototropism. Floral heliotropism

Heliotropic flowers track the sun's motion across the sky from East to West. During the night, the flowers may assume a random orientation, while at dawn they turn again towards the East where the sun rises. The motion is performed by motor cells in a flexible segment just below the flower, called a pulvinus . The motor cells are specialized in pumping potassium ions into nearby tissues changing their turgor pressure. The segment flexes because the motor cells at the shadow side elongate due to a turgor rise. Heliotropism is a response to blue light. Leaf heliotropism Leaf heliotropism is the solar tracking behavior of plant leaves. Some plant species have leaves that orient themselves perpendicularly to the sun's rays in the morning (diaheliotropism), and others have those that orient themselves parallel to these rays at midday (paraheliotropism). Floral heliotropism is not necessarily exhibited by the same plants that exhibit leaf heliotropism. Phototropism in plants

Phototropism is directional growth in which the direction of growth is determined by the direction of the light source. In other words, it is the growth and response to a light stimulus. Phototropism is most often observed in plants, but can also occur in other organisms such as fungi. The cells on the plant that are farthest from the light have a chemical called auxin that reacts when phototropism occurs. This causes the plant to have elongated cells on the farthest side from the light. Phototropism is one of the many plant tropisms. Arabidopsis thaliana is a small flowering plant native to Europe, Asia, and northwestern Africa. A spring annual with a relatively short life cycle, Arabidopsis is popular as a model organism in plant biology and genetics is directed by blue light receptors called phototropin. Other photosensitive receptors in plants include phytochrome. Phytochrome is a photoreceptor, a pigment that plants use to detect light. It is sensitive to light in the red and far-red region of the visible spectrum. Many flowering plants use it to regulate the time of flowering based on the length of day and night and to set circadian rhythms that sense red light and cryptochrome. Phototropism is enabled by auxins. Auxins are plant hormones that have many functions. In this respect, auxins are responsible for expelling protons (by activating proton pumps) which decreases pH in the cells on the dark side of the plant. This acidification of the cell wall region activates enzymes known as expansins. The decrease in cell wall strength causes cells to swell, exerting the mechanical pressure that drives phototropic movement. Other developmental responses of plants to sunlight are flowering in short day and long day plants, etiolation, photonasty, phototaxis and photo orientation. PHOTODESTRUCTIVE EFFECT ON PLANTS

The activity of solar radiation as a stress factor on plants is known as photodestructive effect. Essentially, ionizing radiation are radiations that occurs below 380 nanometers (violet light) on the spectrum. This type of radiation starts with Ultraviolet (UV), then X-ray and down into Gamma rays. Basically what happens is that the radiation breaks apart atomic bonds in molecules which releases electrons and makes ions. That is why it is called ionizing radiation.

Plants are better adapted to cope with naturally occurring ionizing radiation, like UV, than people are because they contain special chemical compounds that protect against UV radiation. Molecules in living material (whether plant or animal) are complex and large in size. Ionizing radiation can cause changes in plants by breaking up these complex molecules. This may change the structure of the plant DNA, creating mutations or it might just kill the cell outright. The loss of a few cells has little effect on the growth of a plant. However, when massive amounts of cells are killed, the plant may become stressed and grow slower or even die. When DNA is mutated, the existing plant may not show any mutations itself but mutations may show up in its offspring. It cannot be predicted what ionizing radiation will do to a plant. Mutations by radiation in plants can easily occur at certain stages. Radiation can change the reproducing parts of the parent plant causing mutations in its offspring. However, in nature, this does not occur often because pollen itself has an extremely tough exterior and contains special pigments that protect against naturally occurring radiation (UV). When people, however, use very high concentrated doses of radiation, these protection mechanisms can be overcome. The next stage where a plant's DNA can be easily mutated is at the seed stage, before the plant begins to grow. And finally, a very young plant is more susceptible to radiation mutations than an older plant.

CONCLUSION With regard to the fact that climatic conditions are always changing, it is a task within the field of plant ecophysiology and paleoecology to investigate and elucidate the causal relationships of the interactions between environmental factors and plant distributions by conducting comparative studies. An important task will be the systemic search for plant species and functional groups that respond to similar environmental influences differently, as they may serve useful bioindicators for changing climatic conditions.


Walter Larcher, Physiological plant ecology: ecophysiology and stress physiology, Springer publications *****