PLANT TROPISMS & HORMONAL CONTROL

At every stage in the life of a plant, sensitivity to the environment and coordination of responses are evident. One part of a plant can send signals to other parts. For example, the terminal bud at the apex of a shoot is able to supress the growth of axillary buds that may be many metres away. Plants keep track of the time of day and the time of year. They can sense gravity and the direction of light. For example, the grass seedling in the photograph above is growing towards light. Plants are constantly tuned to the changing surroundings by interactions between environmental stimuli and internal signals and responses. This section focuses on how plants respond to these external and internal cues. Animals, being mobile, respond mainly by behavioural mechanisms, moving towards positive stimuli and away from negative stimuli. Rooted to one location for life, a plant generally responds to environmental cues by adjusting its pattern of growth and development. For this reason, plants of the same species vary in body form much more than do animals of the same species. All lions have four limbs and approximately the same body proportions, but oak trees vary considerably in their number of limbs and their shapes. tropism: response of plants to stimuli phototropism: response of plant to light geotropism: response of plant to gravity hydrotropism: response of plant to water thigmotropism: response of plant to touch hormone: chemical produced by particular cells, that travel to other cells in body fluids, which affect the functions of other cells capable of responding

How does a hormone pass a signal from one cell to another cell?
Hormones The word hormone is derived from a Greek verb meaning to excite. Found in all multicellular organisms, hormones are chemical signals that coordinate the parts of the organism. By definition, a hormone is a chemical compound that is produced by cells in one part of the body and then transported to other parts of the body, where it binds to a specific receptor and triggers responses in target cells and tissues. Another important characteristic of hormones is that only minute amounts are required to induce substantial change in an organism. Hormone concentrations or the rates of their transport can change in response to environmental stimuli. Often the response of a plant is governed by the interaction of two or more hormones.

Figure 1. Overview of how a hormone stimulates a target cell to respond. A hormone binding to a specific receptor stimulates the cell to produce secondary messengers. Secondary messengers trigger the cells various responses to the original signal. In this diagram, the receptor is on the surface of the target cell. In other cases, hormones enter cells and bind to specific receptors inside.

Response Ultimately, the hormone signal pathway, shown in the diagram above, leads to the regulation of one or more cellular activities. In most cases, especially when changes in development are involved, these responses to stimulation involve the increased activity of certain enzymes. Hormones control the types and rates of chemical reactions in cells, change in movement of substances across the plasma membrane, secretion, growth of cells, and switching a gene on or off. Target cells for a particular hormone are those that possess a specific receptor. Specificity Plant hormones are less specific than animal hormones. Flowering plant hormones tend to affect many or most cells of the plant. While a few hormones affect most cells in an animals body, animal hormones usually affect specific organs, and often only one type of cell within that organ. Plant hormonal plant responses are simpler than animal hormonal responses. Hormoneproducing cells in plants are not organised into an endocrine system of glands. Speed and duration Hormonal effects are generally slower than nervous system responses, but the effects last longer than the effects of nerves. For example, consider a sudden fright. The hormone adrenaline will remain in the body long after the fast neural response has caused the person to jump away or flinch, but the hormonal responses will be slower. Plant hormonal responses are much slower than animal hormonal responses. Moving through a plant hormone pathway is rather slow (xylem and phloem, or from cell to cell to reach target cells) compared with an animal transport system (open or closed circulatory system). Also, the distance that a hormone travels in a plant can be large, which takes time. Consider the length of a long tree branch. Movement of a plant hormone from source cell to target cell requires expenditure of energy stored in ATP molecules. Active transport is about 10 times faster than if the hormone simply moved by diffusion. Even so, movement of the hormone may be as slow as 1 cm per hour, in a plant.

Questions 1. 2. 3. 4. What is a hormone? Why do plant hormones generally move more slowly than animal hormones from source cell to target cell? Which system works fastest? the hormonal system or the nervous system? hormonal system/nervous system Which system affects the body for the longest time? hormonal system/nervous system

Plant responses to hormones Research on how plants grow toward light led to the discovery of plant hormones
The concept of chemical messengers (hormones) in plants emerged from a series of experiments on how stems respond to light. A houseplant on a windowsill grows toward light. If you rotate the plant, it will soon reorient its growth until its leaves again face the window. Any growth response that results in curvatures of parts of a plant toward or away from a stimulus is called a tropism. (from the Greek tropos, turn). The growth of a shoot toward light is called positive phototropism (growth away from light is called negative phototropism). In a forest or other natural ecosystem where plants may be crowded, phototropism directs growing seedlings toward the sunlight that powers photosynthesis. What is the mechanism for this adaptive response? Much of what is known about phototropism has been learned of studies of grass seedlings, particularly oats. The shoot of a grass seedling is enclosed in a covering called the coleoptile, which grows straight upward if the seedling is kept in the dark or if it is illuminated uniformly from all sides. If the growing coleoptile is illuminated with light from one side, it will curve towards the light (see the photograph of the candle that opens this section). This response results from different growth of cells on opposite sides of the coleoptile; the cells on the darker side elongate (lengthen) faster than cells on the brighter side. Charles Darwin and his son conducted some of the earliest experiments on phototropism in the late 19th century. They observed that a grass seedling could bend toward light only if the tip of the coleoptile was present. If the tip was removed, the coleoptile would not curve. The seedling would also fail to grow toward light if the tip was covered with an opaque cap (cap which stops light reaching the coleoptile), nor an opaque shield placed farther down the coleoptile prevented the curving response to light. It was the tip of the coleoptile, the Darwins concluded, that was responsible for sensing light. However, the actual growth response, the curvature of the coleoptile, occurred some distance below the tip. The Darwins postulated that some signal was transmitted downward from the tip of the elongating region of the coleoptile. A few decades later, Peter Boysen-Jensen tested this hypothesis and demonstrated the signal was a mobile chemical substance, or hormone. He separated the tip from the remainder of the coleoptile by a block of gelatin, which would prevent cellular contact but allow chemicals to pass. These seedlings behaved normally, bending toward light. However, if the tip was experimentally segregated from the lower coleoptile by an impermeable barrier, no phototropic response occurred. In 1926, F.W. Went extracted the chemical messenger for phototropism by modifying the experiments of Boysen-Jensen. For this chemical messenger, or hormone, Went choose the name auxin (from the Greek auxein, to increase). Auxin was later purified and its structure determined.

Early experiments of phototropism. Only the tip of the coleoptile can sense the direction of light, but the bending response that occurs some distance below the tip. A signal of some kind must travel downward from the tip. The signal can pass through a permeable barrier (gelatin block) but not through a solid barrier (a mineral called mica), suggesting that the signal for phototropism is a mobile chemical.

The Went experiments. Some chemical (indicated in pink) that can pass into an agar block from a coleoptile tip stimulates elongation of the coleoptile when the block is substituted for a tip. If the block is placed off-center on the top of a decapitated coleoptile kept in the dark, the grass bends as if responding to illumination from one side. The chemical is the hormone auxin, which stimulates elongation of cells in the shoot.

Plant hormones help coordinate growth, development, and response to environmental stimuli
The table below previews some of the major classes of plant hormones; auxin, cytokinins, giberellins, inhibitors (e.g. absicisic acid) and ethylene.

Hormone Auxins

Cytokinins Gibberellins Abscisic acid

Ethylene (ethene)

Major Functions (note: not a complete list) increase the rate of cell elongation, being important in both phototropic and geotropic responses. increase the rate of cell division. promotes apical dominance which occurs when the apex or tip of a plant grows while the lateral buds dont develop. promote cell division (cytokinesis). cause stem elongation by promoting both cell elongation and cell division. inhibits growth. initiates stress responses in plants such as dormancy; closes stomata during water stress. a gas which acts as a hormone and stimulates ripening of fruit and the death of flowers.

Figure .Structure of some plant hormones.

Auxins The term auxin is used for any chemical substance that promotes the elongation of coleoptiles. Although auxin affects several aspects of plant development, one of its chief functions is to stimulate the elongation of cells in young developing shoots.

Cytokinins Trial-and-error attempts to find chemical additives that would enhance the growth and development of plant cells in tissue culture led to the discovery of cytokinins. In the 1940s Johannes van Overbeek found he could stimulate growth of seedlings using coconut milk. The active ingredient of coconut milk turned out to be growth regulators. These growth regulators were named cytokinins because they stimulate cytokinesis, or cell division. It is not known if plants produce their cytokinins or if symbiotic bacteria. Control of cell division and differentiation. Acting in concert with auxin, cytokinins stimulate cell division and influence the differentiation of cells into specialised cells. Control of Apical Dominance. Cytokinins and auxin work together to control apical dominance (the ability of a terminal bud to supress the development of side buds, which causes the plant to grow upright). Cytokinins counter the effects of the auxin, stimulating the growth of axillary buds so that plant grows sideways more and is bushier.

Gibberellins A century ago, farmers in Asia noticed that some rice seedlings in their paddies grew so tall and spindly that they toppled over before they could mature and flower. In 1926, a Japanese plant scientist, discovered that a fungus Gibberella fujikuroi, which produces large amounts of gibberellin, caused this foolish seedling disease. In the 1950s , researchers discovered that plants also make gibberellins. Gibberellins stimulate growth in both the leaves and stem, but not the roots. Gibberellins have a variety of effects in plants which include stem elongation, fruit growth and signalling seeds to break dormancy and germinate.

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Figure 4 Foolish seedling disease in rice. The spindly rice plants on the right are infected with the fungus Gibberella. The pathogen secretes

gibberellins, a growth stimulant that uninfected plants (left) also produce, but in smaller quantity.

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Plant on the left has been treated with gibberellin. The plant on the right is the control.

Abscisic Acid In the 1960s , one research group studying the chemical changes preceding leaf abscission (the dropping of autumn leaves), isolated the hormone abscisic acid (ABA). Ironically, although named from the word abscission, abscisic acid is no longer thought to play a primary role in leaf abscission, but is a plant hormone of great importance in other functions. Unlike the growth-stimulating hormones we have studied so far auxin, cytokinins, and gibberellins abscisic acid generally slows down growth. Often ABS antagonizes (opposes) the actions of the growth hormones, and it is the ratio of ABA to one or more growth hormones that determines the final growth outcome. ABA has many effects on plants, but we will consider just two.
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Seed Dormancy. Seed dormancy has great survival value because it ensures that the seed will only germinate when there are optimal conditions of light, temperature, and moisture. What prevents a seed dispersed in autumn from germinating immediately only to be killed by winter? What mechanisms ensure that such seeds germinate in the spring? For that matter, what prevents seeds from germinating in the dark, moist interior of the fruit? The answer to these questions is ABA. The high levels of ABA in maturing seeds inhibit germination and induce the production of special proteins that help the seeds withstand the extreme dehydration that accompanies maturation. Many types of dormant seeds will germinate when ABA is removed or inactivated in some way. The seeds of some desert plants break dormancy only when heavy rains wash ABA out of the seed. Other seeds require light or prolonged exposure to cold to trigger the inactivation of ABA. Drought stress. ABA is the primary internal signal that enables plants to withstand drought. When a plant begins to wilt, ABA accumulates in the leaves and causes stomata to close rapidly, reducing transpiration and preventing further water loss. ABA causes a rapid loss of potassium from guard cells. The accompanying osmotic loss of water leads to a reduction in guard cell turgor (pressure) and a closing of the stomata (see diagram below).

Ethylene During the 19th century when coal gas was used for street illumination, the leakage of illuminating gas from gas mains caused nearby leaves to drop their leaves prematurely. In 1901 a Russian scientist named Dimitry Neljubow demonstrated that the gas ethylene was the active factor in illuminating gas. The idea that ethylene is a hormone that plants produce only became accepted, however, when the technique of gas chromatography became available. Plants produce ethylene (also known as ethene) in response to stresses such as drought, flooding, mechanical pressure, injury, and infection. Ethylene has numerous effects on plants, but we shall focus on just four.

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The Triple Response to Mechanical Stress: Imagine a pea seedling pushing upward through the soil only to come up against the underside of an immovable object such as a stone. As the stem pushes against the obstacle, the mechanical stress in its delicate tip induces the seedling to start producing ethylene. Ethylene, in turn, instigates the seedling to perform a growth manoeuvre called the triple response that enables it to circumvent the object. The three parts of this response, which you can see in the figure below are a slowing of stem elongation, a thickening of the stem (which makes it stronger), and a curvature that causes the stem to start growing horizontally. As the stem continues to grow, its tip touches upward intermittently. If these probings continue to detect a solid object above , then another pulse of ethylene is generated and the stem continues its horizontal progress. If, however, the upward touch detects no solid object, then ethylene production decreases, and the stem, now clear of the obstacle, resumes its normal upward growth. This is how plants grow around any obstacles as they grow upwards.

Figure 4 Ethylene induces the triple response in pea seedlings to grow around obstacles.

Either artificial treatment with the gaseous hormone ethylene or natural production of the hormone triggered by mechanical stress causes the stems of germinating pea seedlings to elongate less rapidly, to become thicker, and to grow horizontally the triple response. The growth response is an adaption that helps the seedling circumvent obstacles encountered during soil penetration. This experiment shows the extent of the triple response in peas seedlings treated with different concentrations of ethylene.

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Apoptosis: Programmed Cell Death. Consider the shedding of a leaf in autumn or the death of an annual (plant that lives one growing season only) after flowering. Or think about the final step in the differentiation of a xylem vessel, when its living contents are destroyed, leaving a hollow transport tube behind. The cells, organs, and plants that are genetically programmed to die on a particular schedule do not simply shut down their cellular machinery and await death. Rather, the onset of programmed cell death, called apoptosis, is one of the busiest times in a cells life, requiring new gene expression.

Autumn Melody photograph by Tom Wundrak Hormones control when deciduous leaves fall.

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