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S E C T I O N 15.1 Plant Responses to Stimuli E X P E C TAT I O N S Describe the effects of growth regulators. Design and carry out experiments to study factors that affect the growth of plants. Describe the structure and function of a vascular plant. Describe some of the industrial processes that use plants. Figure 15.1 Some plants can propogate, or grow new individuals, from a leaf. Plants as living organisms have the ability to: use energy to obtain materials from the external environment, and use energy to rearrange those materials into new plant substances. With very few exceptions, plants acquire all the matter and energy they need: without changing location (as must most animals, protists, and bacteria), without preying on other living organisms (as animals do), and without relying on matter assembled by other organisms (as fungi do). Some plants, such as the Bryophyllum shown in Figure 15.1 can grow new individuals from a leaf. Some trees, such as sumac and poplar, grow new individuals, called suckers, from the roots. Spider plants and strawberries grow stems from which new individuals can become established at some distance from the original plant. This is shown in Figure 15.2. Many grasses grow from nodes in their roots. A field that has been ploughed or overgrazed will be covered by a new blanket of grasses that grew from the roots of the old plants. Wo rd LINK The term “plant” comes from the Latin word planta which means sprout, slip, or cutting. Today we apply the term “plant” to entire organisms, not just slips or cuttings. 554 MHR • Plants: Anatomy, Growth, and Functions Figure 15.2 Plants such as this spider plant are able to grow new individuals from stems from the plant. Herbivorous animals eat and gain nutrients from plants. Most plants however, obtain their nutrients from the air and the soil. Some plant species however, are able to “prey”, or eat and gain nutrients from animals. Figure 15.3 on the next page shows the pitcher plant (the provincial flower A pitcher plant B Venus’s-flytrap — open C Venus’s-flytrap — closed Figure 15.3 Plants such as the pitcher plant and the Venus’s-flytrap are consumers. They are not able to meet their nutritional requirements from the air and the soil. of Newfoundland and Labrador) and the Venus’sflytrap, both of which are consumers. They trap and digest insects in order to obtain nutrients that are not available in the nutrient-poor soil they grow in. These plants are also producers because they photosynthesize, as well. repeatedly, they begin to differ in shape, in the relative proportions of their various organelles, and in the functions they can perform. These changes result in the cells becoming specialized for particular functions, such as photosynthesis, storage, and support. Development in the Meristem What gives plants their amazing ability to grow throughout their lives? Although mitosis and cell division occurs throughout a plant as it grows, eventually new growth is restricted to small regions of unspecialized tissue collectively called the meristems shown in Figure 15.4. Growth in meristem tissue results from the accumulation of rapidly dividing cells. When a cell in the meristem divides, one of the two resulting cells remains in the meristem. The other cell becomes part of the plant body. Initially, all meristem cells are identical in structure, and they have no specialized function. As they divide terminal meristem shoot apical meristem lateral meristems root apical meristems Figure 15.4 At first, the small, actively dividing cells of meristem tissue are identical. With repeated division, they “differentiate.” That is, they become specialized for different plant functions. Using Plants • MHR 555 Types of Meristem Tissue annual growth rings vascular cambium Figure 15.5 The meristem in a tree stem is located in the vascular cambium. DESIGN YOUR OWN Investigation There are two main types of meristem tissue. The first type is called apical meristem tissue. It is located in the root and shoot tips of plants. Division of apical meristem cells results in growth of roots, leaves, and flowers. In the root, the apical meristem is protected by a root cap. In the stem it is protected within a terminal bud. In colder climates, the terminal buds stop growing in the winter and resume growing in the spring. These buds are protected by bud scales, which fall off when growth begins in the spring. Lateral meristem results in the growth of tissue beneath the bark of tree stems, as shown in Figure 15.5. The division of cells in the lateral meristems results in the thickening of cylinders of tissue. Most woody plants have two kinds of lateral meristems: a vascular cambium and a cork cambium. SKILL FOCUS 1 5 • A Hypothesizing Factors Affecting Plant Growth Identifying variables There are many different factors that will determine how well a plant will grow. What external factors affect plant growth? The list includes: the amount of light; the type of soil; the amount of water; the amount of minerals and other nutrients; and temperature. Which of these factors can be explored in an experimental setting? In this investigation, you will design an experiment to identify and investigate one external factor affecting plant growth. Problem What is the effect of an external factor on the growth of plants? Make a hypothesis about the effect of an external factor (of your choice) on plant growth. CAUTION: Wash your hands thoroughly after working with soil. potting soil (or other factors that you will be types of potting material) testing such as different fertilizers, water, or salt 1. Decide which condition you are going to vary in your experiment. Prepare a list of possible ways you might test your hypothesis. 2. Decide on one approach for your experiment that could be conducted in the classroom. 3. Your design should test one variable at a time. Plan to collect quantitative data. Materials 556 Analyzing and interpreting Experimental Plan Hypothesis bean seeds (or other easy to grow seeds) labels Performing and recording plant pots or planting trays water ruler MHR • Plants: Anatomy, Growth, and Functions 4. Outline a procedure for your experiment listing each step. The lateral meristem, called the vascular cambium produces xylem and phloem cells in the stems and roots. The other lateral meristem, called the cork cambium, produces a tough covering for the surface of stems and roots. The outer bark of a tree is produced by the cork cambium. Meristem tissue enables plants to grow from cuttings. Growing plants from cuttings is the basis of plant cloning. Cloning technology, especially relevant to agriculture, is the process of growing genetically identical copies of an organism from a single cell or part of an organism. For some species, growing plants from cuttings can be much faster than growing them from seed. REWIND Turn to Chapter 6, Section 6.2, to review cloning techniques and the results of cloning experiments. 5. Show an outline of your plan to your teacher. Get your teacher’s approval before continuing. Checking the Plan PAUSE RECORD Make a chart with two columns headed “now” and “later.” In the “now” column, record a definition of plant as you understand the term today, using your knowledge from Chapters 13 and 14. Your personal definition will grow and develop as you progress through this chapter. Use the “later” column to record your new understandings. On a separate page, write your ideas of why plants are important to Earth, the weather, animals, and other plants. Internal Regulation of Plant Growth and Development Plants can grow to their maximum height when environmental conditions are optimal. Optimal conditions include adequate moisture, warmth, light, and nutrients. Fertilizers promote plant growth and development by providing additional nutrition. Pesticides and fungicides promote plant 2. Suggest reasons why your plant may not have grown as well in the other conditions. 3. Do your results support your hypothesis? Give reasons for your answer. 1. What will be your independent variable? What will be your dependent variable(s)? What will be your control variable(s)? 4. If the results were different than what you expected, what might be the reasons for these results? 2. What will your experimental steps be? How would you apply the different treatments to the plants in the appropriate labelled pots? 5. Determine which treatment was the most beneficial and which treatments were the least beneficial. Look at other factors that may have affected your experiment. Were there any that you did not control? 3. What will you measure or count? How will you determine if your variable has had an effect on plant growth? 4. Design a table to record your data. Data and Observations Conduct your experiment and make your measurements. Enter the data into your table. Make a graph of your results. CAUTION: Make sure you are wearing all the appropriate safety equipment while you perform your experiment. Analyze 1. At what level of the variable you were testing did your plant grow best? Conclude and Apply 6. Based on your results, what recommendations would you make to farmers growing crops of your plant? Exploring Further 7. What would happen if you varied two of the treatments at one time? Would you be able to tell which treatment affected the plants? 8. How do scientists determine what factors affect plant growth under field conditions when there are many variables (or treatments) that are affecting growth? Using Plants • MHR 557 growth by controlling numbers of insects and fungi that feed on plants. However, plant growth and development are also controlled by the plant’s own hormones. From your previous studies, you learned that a hormone is a chemical compound manufactured by specialized tissue in one body part of an organism but that governs or regulates the activity of another body part or parts. Even though a hormone may circulate throughout an organism, it will act only on specific “target” tissues or organs. Investigation Hormonal Control of Plant Growth In the early 1800s, experiments undertaken by Charles Darwin and his son Francis described the effects of a mysterious “influence” that affected the growth of grass seedlings. The seedlings normally grew toward a light source, however this behaviour was not seen if the tips of the grass seedlings were covered with an opaque capsule that did not let light through. The remainder of the plant, where the growth actually occurs, was still exposed to light. If the tip of the seedling were covered with a gelatin capsule, which allowed light to pass through, then the seedling would grow towards the light as expected. For many years, researchers SKILL FOCUS 1 5 • B Predicting How Gibberellic Acid Affects Plant Growth and Development Performing and recording Many garden centres stock chemicals that promote growth. Brand names differ, but the labels reveal that many contain the same main ingredient: gibberellic acid. What does this compound do for plants? Conducting research Pre-lab Questions Procedure What are the classes of plant hormone growth promoters? What are the effects of these different classes of hormones? Problem How does gibberellic acid affect the growth of bean plants? Prediction Predict what will happen to the rate of growth as plants absorb gibberellic acid. CAUTION: Wash your hands thoroughly after handling seeds, seedlings, soil, or gibberellic acid. Materials recycled 6-compartment planting tray labels and marking pens soil or potting mixture water 558 Analyzing and interpreting pre-soaked bean seeds, 6 per group ruler gibberellic acid graph paper MHR • Plants: Anatomy, Growth, and Functions 1. Label the tray with your group’s name. Fill each compartment with soil or potting mixture. 2. Plant one pre-soaked bean seed in each compartment. Place the tray in a brightly lit location that is warm enough for plant growth but not too hot. 3. Every day, add enough water to keep the potting mixture moist, using equal amounts of water in each compartment. 4. When most seedlings are over 10 cm tall, select three seedlings that are equivalent in height. Mark them and leave them in the tray. Remove the other three. This is Day 1 of your investigation. 5. Join another group so that two trays are available. Prepare two labels, one for each tray: “GA YES” (meaning treat with gibberellic acid) and “GA NO” (meaning do not treat with gibberellic acid). Prepare an observation chart in your notebook like the one shown on the next page. conducted experiments to try to explain the nature of this observation. In 1926, Frits Went performed a series of experiments that showed there was a chemical messenger in the grass seedlings. This chemical could enhance plant growth. He named the substance auxin, from the Greek work auxein, which means “to increase”. Other discoveries about plant hormones came as a result of people noticing unusual growth in plants. For example, observers noticed that a rice plant infected with the fungus Gibberella fujikoroi grew abnormally tall. In 1935, researchers were finally able to isolate the chemical compound that caused the accelerated growth, and named it Average height of seedlings in cm over several days Tray Day 1 Day 2 Day 3 Day 4 Day 5 GA No GA Yes 6. Measure the height of each individual seedling. Calculate the average height for the seedlings in each tray and record your results in the Day 1 column. 7. Treat the plants in the “GA YES” tray by placing one drop of gibberellic acid on the shoot tip of each seedling. 8. Repeat Steps 6 and 7 for up to 5 days, recording your results in the appropriate column. Be sure all seedlings receive the same amount of light, warmth, and water each day. 9. Watch for any obvious difference in growth between the two trays. Note when this difference becomes apparent. gibberellic acid. They discovered that applying artificially-manufactured gibberellic acid to a plant not infected by the fungus caused the plant to grow abnormally tall. Scientists and researchers continued to search for other plant hormones that might affect growth and development. They discovered two types of plant growth hormones: promoter hormones, which are hormones that cause growth, and inhibitor hormones, which are hormones that block growth. 2. Based on your observations, how does gibberellic acid affect the growth of the bean seedlings? How soon after the first application can this effect be observed? 3. What would you expect to observe if the gibberellic acid were applied to the base of the stem rather than to the tip of the shoot? Conclude and Apply 4. What medium did your plants grow in? Where did you place the drops of gibberellic acid? Do you think gibberellic acid would make a good additive for impoverished soil? Explain why or why not. 5. Look at this sketch of a normal rice plant. Sketch the probable appearance of a rice plant infected with the fungus Gibberella fujikuroi. How would you know the infection is present? 10. Use the data in your observation chart to create a graph. Plot height vs. time for both trays on the same piece of graph paper. Label the curves appropriately. Post-lab Questions 1. What variable were you testing? Which tray was the experimental group? Which tray was the control group? What variables were you controlling in both trays? Using Plants • MHR 559 Table 15.1 Naturally occurring plant growth hormones I Class Auxins II Example III Molecular structure β -indoleacetic acid CH 2 COOH N H Cytokinins H 6-furfurylaminopurine CH 2 N Figure 15.6 As auxin concentrations decrease, the deciduous trees begin to shed their leaves. The Classes of Hormones The different classes of plant hormones and their molecular structure are summarized in Table 15.1. A full summary of the type and function of all of the plant hormones is given in Table 15.2 on the next page. Auxins are a class of hormone that is produced in the apical meristem of shoots. There are both natural and synthetic auxins that promote cell elongation, the development of vascular tissue, and trigger the development of above-ground stems, which help support the plant. These hormones also cause leaves to drop after they are no longer needed. The stimulation of cell elongation occurs because auxin increases the plasticity of the plant cell wall. The more plastic the cell wall is, the more it can stretch during active cell growth. This process is shown in Figure 15.7. N N H Gibberellins Gibberellic acid (GA3) O HO C O OH CH 2 CH 3 Abscisic acid COOH Abscisic acid (ABA) CH 3 CH 3 O Ethylene Ethylene gas CH 3 CH C OH CH CH CH 3 COOH H H C H cell wall H+ ATP ATP H+ O N N C H Note: Three of the hormone classes listed in column I are shown as plurals. Each of these classes includes two or more chemical compounds with similar functions but different structures or formulas. Only the most common example of each is shown in column II. ATP H+ ATP auxin ATP H+ Figure 15.7 Auxin stimulates an H+ pump so that hydrogen ions H+ 560 MHR • Plants: Anatomy, Growth, and Functions (H+ ) are transported out of the cytoplasm of the cell. The resulting acidity causes the cell wall to weaken, and solutes enter the cell. Water follows by osmosis and the cell elongates. Gibberellins are also produced in the apical meristem and act to increase stem length. As well, gibberellins increase the uptake of starch in the embryo of germinating seeds and stimulate the development of vascular tissue. The effects of gibberellins include taller and stronger plants, plants that flower early, or genetically dwarf plants that grow to normal heights. Gibberellins are used in commercial crops to increase fruit size, and cluster size in grapes. As well, they can delay the ripening of citrus fruits and speed up the flowering of strawberries. Cytokinins promote cell division and cell differentiation. Cell differentiation occurs when specialized cells are needed to perform certain functions. They also delay the aging of leaves and fruit. Cytokinins work by influencing the synthesis and activation of proteins that are required for mitosis. Oligosaccharins are a recently discovered class of growth promoters. They stimulate plants to manufacture antibiotics in response to attack by fungi or bacteria. This allows the plant to grow to its full potential because the negative influences of pests are diminished. In addition to hormones that promote plant growth, plants produce hormones that inhibit growth. There are two classes of hormone growth inhibitors. The first is abscisic acid (ABA), which is synthesized mainly in mature green leaves, fruits, and root caps. This hormone inhibits the germination of seeds, inhibits the growth of buds in plant stems, and blocks the intake of carbon dioxide by controlling the opening and closing of leaf stomata. Abscisic acid also blocks the action of growth promoting hormones. Ethylene, the second inhibitor, is a gaseous hydrocarbon. It occurs as a natural plant hormone. It stimulates the aging of plant tissues, the ripening and sweetening of fruit, and can also speed up the dropping of leaves from trees. The production of ethylene gas by plants can stimulate other plants to ripen. This was initially noted when bananas ripened quickly if they were left near oranges. The ripening action of ethylene has led to its use in agriculture. For example, tomatoes may be picked while green and then ripened artifically by the application of ethylene. Table 15.2 Summary of plant growth hormones and their actions Growth promoters Auxins act to promote cell elongation suppress the growth of lateral branches trigger the growth of prop roots that grow from aboveground stems suppress leaf drop before leaf ages; promote leaf drop afterward Gibberellins act to promote cell enlargement promote uptake of starch tissue by the embryos in germinating seeds reverse genetic dwarfism stimulate the vascular cambium to produce secondary phloem in woody plants promote “bolting,” the rapid elongation of the flower stem in plants such as cabbage Cytokinins act to promote cell division (recall that cytokinesis is the process of cell division after mitosis) stimulate formation of adventitious buds delay senescence (ageing) of leaves by maintaining chlorophyll content reverse suppression by auxin Growth inhibitors Abscisic acid (ABA) acts to block intake of carbon dioxide by causing the closure of leaf stomata inhibit seed germination inhibit active growth of axillary lateral buds block the action of growth-promoting hormones promote abscission of leaves and fruits Ethylene acts to stimulate fruits to ripen stimulate other effects associated with tissue ageing trigger its own production through positive feedback (its presence promotes the production of still more ethylene, so ethylene levels build up rapidly) Using Plants • MHR 561 Plant Tropisms Plants exhibit the ability to orient themselves in response to external stimuli such as light. A directional growth response to unequal stimulation from the external environment is called a tropism, and it controls the growth pattern of the plant. Various external stimuli affect the production of plant hormones. This results in the directional growth of a plant. In tropism, the plant may grow either toward or away from the stimulus. Growth toward the stimulus is a positive tropism. Growth away from the stimulus is a negative tropism. There are three major kinds of plant tropisms that are affected by light, gravity, and touch. Phototropism occurs when the growth of a plant is affected by light. In general, plants are positively phototropic, that is, they grow toward light. Roots are negatively phototropic and grow away from light. The growth is caused by differing amounts of auxin produced on the light and dark sides of the stem. Auxin accumulates on the shaded side of the stem, which causes the cells there to elongate. This causes the stem bend toward the light, as shown in Figure 15.8. Turning the plant around will cause the stem to bend in the other direction however, it will not change the original curve in the stem because it is the result of growth. Figure 15.9 demonstrates the experimental procedure that led to the discovery of phototropism. light source elongated cells Figure 15.8 Tropisms result when external stimulation is unequal. For example, the stem of this plant is receiving much more light on its right surface than on its left surface. Figure 15.9 Oat seedlings, After tips are placed on agar, agar is cut into blocks. Coleoptile tip is intact. MINI Coleoptile tip is removed. Cell elongation occurs only beneath block. LAB How Plants Respond to Light The flowering heads of sunflowers can be seen to turn in response to the movement of the Sun as it moves across the sky. This is one example of phototropism. Plant some bean, tomato, or other vegetable seeds or acquire some seedlings. When the plants are a few inches high, place them in a room where the only light comes from one light source. (For example, place them on a window ledge with a cardboard box blocking out the light from the classroom.) Examine the plants daily and record what you see. 562 Block is placed to one side of coleoptile. shown here, are protected by a hollow sheath called a coleoptile. After a tip is removed and placed on a block of agar, a block of that agar placed on one side of the coleoptile can cause it to curve even in the absence of light. The agar blocks contain the hormone produced by the original coleoptile. MHR • Plants: Anatomy, Growth, and Functions Analyze 1. How long does it take to see any bending of the stems? 2. How could you get the plant to straighten out? 3. Where would you expect the auxin to be produced in these plants? Gravitropism is a plant’s response to gravity. This tropism causes roots to grow downwards (positive gravitropism) and shoots and stems to grow upward (negative gravitropism). This benefits the plant, because shoots that grow upward will receive light and roots that grow downward will receive nutrients from the soil. Thigmotropism is the response of plants to touch. This behaviour is a caused by specialized cells in the epidermis of the plant. Vining plants demonstrate a strong positive thigmatropism, as shown in Figure 15.10. The vines grow toward the object touching them causing them to coil around the object. Other plants demonstrate a negative thigmotropism. Figure 15.10 Vining plants such as bindweed demonstrate thigmotropism. PAUSE RECORD Record how you would expect the primary root of a germinating seed to respond to light and to gravity. Name these tropisms. Repeat for the stem of a germinating seed. Write a brief explanation of why you would expect these responses. Nastic Responses in Plants Another type of response, called nastic movements, are caused by a stimulus that is not directional. For example, the leaves on a mimosa plant fold up when the plant is touched, as shown in Figure 15.11. This response might seem to illustrate a negative thigmotropism, however, it is neither directional nor permanent. The leaflets fold downward in the same way regardless of the direction of the stimulus. These movements are not a result of growth, but rather a change in turgor pressure in the cells at the base of each leaflet. A sudden drop in pressure causes the cells to become limp and the leaflets fold down. Once the stimulus has ceased, the turgor pressure in the cells rises once again and the leaflets open. Another example of a nastic response is shown in Figure 15.3 on page 555, where you can see the hinged leaf of a Venus’sflytrap. The movement of an insect on the leaf triggers the hinged leaf to close, trapping the insect between the leaves. Commercial Use of Growth Regulators Over the past century, scientists have learned much about plant growth hormones. Horticulturists and other agricultural scientists use this knowledge of plant growth regulators to influence the growth and development of crops and ornamental plants. Most growth regulating hormones used for commercial purposes are synthetically produced rather than extracted from plants. For example, there appears to be only one naturally occuring auxin, but many more synthetic auxin-like growth regulators exist. Although these synthetically produced hormones are not identical to natural auxins, their chemical action is similar, and the plants respond as they would to naturally occurring auxin. Commercial uses of several types of plant growth regulators are listed in Table 15.3 on the next page. Figure 15.11 Mimosa is sometimes called the sensitive plant. When it is touched, it folds its leaves in less than one-tenth of a second. Using Plants • MHR 563 Table 15.3 Some uses for commercial plant growth regulators Plant growth regulators Auxin-like growth regulators Commercial uses stimulate rooting act as a herbicide for dicots prevent sprouts in pruned trees prevent fruit from dropping too soon Cytokinin-like regulators promote axillary bud growth in orchids and daylilies (results in more flowers) prevent browning in cut salads increase fruit size Gibberellic acid-like regulators increase flower size (for example, camellias) increase grape size stimulate separation of berries from stalks stimulate seed germination Gibberellic acid-like inhibitors control (restrict) height in flowerpot plants (lilies, orchids) control height of bedding plants Ethylene-like regulators stimulate flowering stimulate ripening (bananas, tomatoes) accelerate colour development in tomatoes and citrus fruits stimulate separation of cherry stems from branches (for mechanical harvesting machines) Ethylene inhibitors allow long-term, controlled storage of apples allow hypobaric (low-pressure) storage of many fruits, vegetables, and flowers 564 MHR • Plants: Anatomy, Growth, and Functions A large industry is based on the manufacture of artificial plant growth regulating hormones. Some of the growth regulating hormones that are produced can be very specialized For example, a chemical treatment can be applied to ornamental trees to prevent them from growing too tall and interfering with utility lines. From the ground, the trees look normal, but the tops look as if they have been pruned flat. Table 15.3 outlines more of the commercial uses of plant growth regulators. Chemical Look-alikes: Commercial Growth Regulators Over the past century, scientists have built up an impressive body of knowledge about plant growth hormones. As in any branch of science, there is more to be learned and research is ongoing. However, horticulturists are already taking advantage of our present knowledge by applying plant growth regulators to influence the growth and development of crops and ornamental plants. PAUSE RECORD What have you learned about growth regulators? Do you think there should be more public awareness of these chemicals? Suggest how public awareness of plant growth technology could be improved. List two reasons to support your position. SECTION REVIEW 1. K/U Both plants and animals depend on hormones to regulate growth and development. Name three of these hormones and describe their effects. 2. K/U Which plant growth hormone is gaseous? What is its function? 3. C How does a plant growth regulator differ from a plant growth hormone? Explain what the two have in common. 4. I Your uncle buys two unripe pears. He puts the first in a plastic bag by itself and puts the second in a plastic bag with a ripe banana. The second pear becomes edible much faster than the first. Develop a hypothesis to explain why this happens. Give reasons for your answer. 5. K/U Which of the following effects would you expect to observe if the tip of a shoot is cut off? Support your answer. The plant will become taller because the production of gibberellins will be increased. 6. C Where is meristem tissue located? Explain how meristem tissue works. Draw a labelled diagram of apical or lateral meristem tissue to support your answer. 7. MC Plant growth regulators are involved in the cultivation, processing, and storage of many commercially grown fruits and vegetables. Yet, few shoppers are aware of this. Should supermarkets post signs identifying produce treated with plant growth regulators? Give reasons for your answer. 8. I A researcher is investigating how an auxincytokinin mixture affects the development of meristematic tissue. When the mixture contains a higher concentration of auxin, the meristematic tissue develops into organized root tissue. When the cytokinen is more concentrated, the meristematic tissue develops buds. What would you expect to observe if the mixture contained equal mixtures of the two hormones? Explain your reasoning. The leaves will fall off because the tip will release ethylene. Using Plants • MHR 565