Identify common sensory systems and responses in plants
Animals can respond to environmental factors by moving to a new location. Plants, however, are rooted in place and must respond to the surrounding environmental factors. Plants have sophisticated systems to detect and respond to light, gravity, temperature, and physical touch. Receptors sense environmental factors and relay the information to effector systems—often through intermediate chemical messengers—to bring about plant responses.
- Describe how red and blue light affect plant growth and metabolic activities
- Discuss gravitropism
- Describe thigmotropism, thigmonastism, and thigmogenesis
- Explain how plants defend themselves from predators and respond to wounds
Plant Responses to Light
Plants have a number of sophisticated uses for light that go far beyond their ability to photosynthesize low-molecular-weight sugars using only carbon dioxide, light, and water. Photomorphogenesis is the growth and development of plants in response to light. It allows plants to optimize their use of light and space. Photoperiodism is the ability to use light to track time. Plants can tell the time of day and time of year by sensing and using various wavelengths of sunlight. Phototropism is a directional response that allows plants to grow towards, or even away from, light.
The sensing of light in the environment is important to plants; it can be crucial for competition and survival. The response of plants to light is mediated by different photoreceptors, which are comprised of a protein covalently bonded to a light-absorbing pigment called a chromophore. Together, the two are called a chromoprotein.
The red/far-red and violet-blue regions of the visible light spectrum trigger structural development in plants. Sensory photoreceptors absorb light in these particular regions of the visible light spectrum because of the quality of light available in the daylight spectrum. In terrestrial habitats, light absorption by chlorophylls peaks in the blue and red regions of the spectrum. As light filters through the canopy and the blue and red wavelengths are absorbed, the spectrum shifts to the far-red end, shifting the plant community to those plants better adapted to respond to far-red light. Blue-light receptors allow plants to gauge the direction and abundance of sunlight, which is rich in blue–green emissions. Water absorbs red light, which makes the detection of blue light essential for algae and aquatic plants.
The Phytochrome System and the Red/Far-Red Response
The phytochromes are a family of chromoproteins with a linear tetrapyrrole chromophore, similar to the ringed tetrapyrrole light-absorbing head group of chlorophyll. Phytochromes have two photo-interconvertible forms: Pr and Pfr. Pr absorbs red light (~667 nm) and is immediately converted to Pfr. Pfr absorbs far-red light (~730 nm) and is quickly converted back to Pr. Absorption of red or far-red light causes a massive change to the shape of the chromophore, altering the conformation and activity of the phytochrome protein to which it is bound. Pfr is the physiologically active form of the protein; therefore, exposure to red light yields physiological activity. Exposure to far-red light inhibits phytochrome activity. Together, the two forms represent the phytochrome system (Figure 1).
The phytochrome system acts as a biological light switch. It monitors the level, intensity, duration, and color of environmental light. The effect of red light is reversible by immediately shining far-red light on the sample, which converts the chromoprotein to the inactive Pr form. Additionally, Pfr can slowly revert to Pr in the dark, or break down over time. In all instances, the physiological response induced by red light is reversed. The active form of phytochrome (Pfr) can directly activate other molecules in the cytoplasm, or it can be trafficked to the nucleus, where it directly activates or represses specific gene expression.
Once the phytochrome system evolved, plants adapted it to serve a variety of needs. Unfiltered, full sunlight contains much more red light than far-red light. Because chlorophyll absorbs strongly in the red region of the visible spectrum, but not in the far-red region, any plant in the shade of another plant on the forest floor will be exposed to red-depleted, far-red-enriched light. The preponderance of far-red light converts phytochrome in the shaded leaves to the Pr (inactive) form, slowing growth. The nearest non-shaded (or even less-shaded) areas on the forest floor have more red light; leaves exposed to these areas sense the red light, which activates the Pfr form and induces growth. In short, plant shoots use the phytochrome system to grow away from shade and towards light. Because competition for light is so fierce in a dense plant community, the evolutionary advantages of the phytochrome system are obvious.
In seeds, the phytochrome system is not used to determine direction and quality of light (shaded versus unshaded). Instead, is it used merely to determine if there is any light at all. This is especially important in species with very small seeds, such as lettuce. Because of their size, lettuce seeds have few food reserves. Their seedlings cannot grow for long before they run out of fuel. If they germinated even a centimeter under the soil surface, the seedling would never make it into the sunlight and would die. In the dark, phytochrome is in the Pr (inactive form) and the seed will not germinate; it will only germinate if exposed to light at the surface of the soil. Upon exposure to light, Pr is converted to Pfr and germination proceeds.
Plants also use the phytochrome system to sense the change of season. Photoperiodism is a biological response to the timing and duration of day and night. It controls flowering, setting of winter buds, and vegetative growth. Detection of seasonal changes is crucial to plant survival. Although temperature and light intensity influence plant growth, they are not reliable indicators of season because they may vary from one year to the next. Day length is a better indicator of the time of year.
As stated above, unfiltered sunlight is rich in red light but deficient in far-red light. Therefore, at dawn, all the phytochrome molecules in a leaf quickly convert to the active Pfr form, and remain in that form until sunset. In the dark, the Pfr form takes hours to slowly revert back to the Pr form. If the night is long (as in winter), all of the Pfr form reverts. If the night is short (as in summer), a considerable amount of Pfr may remain at sunrise. By sensing the Pr/Pfr ratio at dawn, a plant can determine the length of the day/night cycle. In addition, leaves retain that information for several days, allowing a comparison between the length of the previous night and the preceding several nights. Shorter nights indicate springtime to the plant; when the nights become longer, autumn is approaching. This information, along with sensing temperature and water availability, allows plants to determine the time of the year and adjust their physiology accordingly. Short-day (long-night) plants use this information to flower in the late summer and early fall, when nights exceed a critical length (often eight or fewer hours). Long-day (short-night) plants flower during the spring, when darkness is less than a critical length (often eight to 15 hours). Not all plants use the phytochrome system in this way. Flowering in day-neutral plants is not regulated by daylength.
The word “horticulturist” comes from the Latin words for garden (hortus) and culture (cultura). This career has been revolutionized by progress made in the understanding of plant responses to environmental stimuli. Growers of crops, fruit, vegetables, and flowers were previously constrained by having to time their sowing and harvesting according to the season. Now, horticulturists can manipulate plants to increase leaf, flower, or fruit production by understanding how environmental factors affect plant growth and development.
Greenhouse management is an essential component of a horticulturist’s education. To lengthen the night, plants are covered with a blackout shade cloth. Long-day plants are irradiated with red light in winter to promote early flowering. For example, fluorescent (cool white) light high in blue wavelengths encourages leafy growth and is excellent for starting seedlings. Incandescent lamps (standard light bulbs) are rich in red light, and promote flowering in some plants. The timing of fruit ripening can be increased or delayed by applying plant hormones. Recently, considerable progress has been made in the development of plant breeds that are suited to different climates and resistant to pests and transportation damage. Both crop yield and quality have increased as a result of practical applications of the knowledge of plant responses to external stimuli and hormones.
Horticulturists find employment in private and governmental laboratories, greenhouses, botanical gardens, and in the production or research fields. They improve crops by applying their knowledge of genetics and plant physiology. To prepare for a horticulture career, students take classes in botany, plant physiology, plant pathology, landscape design, and plant breeding. To complement these traditional courses, horticulture majors add studies in economics, business, computer science, and communications.
The Blue Light Responses
Phototropism—the directional bending of a plant toward or away from a light source—is a response to blue wavelengths of light. Positive phototropism is growth towards a light source (Figure 2), while negative phototropism (also called skototropism) is growth away from light.
The aptly-named phototropins are protein-based receptors responsible for mediating the phototropic response. Like all plant photoreceptors, phototropins consist of a protein portion and a light-absorbing portion, called the chromophore. In phototropins, the chromophore is a covalently-bound molecule of flavin; hence, phototropins belong to a class of proteins called flavoproteins.
Other responses under the control of phototropins are leaf opening and closing, chloroplast movement, and the opening of stomata. However, of all responses controlled by phototropins, phototropism has been studied the longest and is the best understood.
In their 1880 treatise The Power of Movements in Plants, Charles Darwin and his son Francis first described phototropism as the bending of seedlings toward light. Darwin observed that light was perceived by the tip of the plant (the apical meristem), but that the response (bending) took place in a different part of the plant. They concluded that the signal had to travel from the apical meristem to the base of the plant.
In 1913, Peter Boysen-Jensen demonstrated that a chemical signal produced in the plant tip was responsible for the bending at the base. He cut off the tip of a seedling, covered the cut section with a layer of gelatin, and then replaced the tip. The seedling bent toward the light when illuminated. However, when impermeable mica flakes were inserted between the tip and the cut base, the seedling did not bend. A refinement of the experiment showed that the signal traveled on the shaded side of the seedling. When the mica plate was inserted on the illuminated side, the plant did bend towards the light. Therefore, the chemical signal was a growth stimulant because the phototropic response involved faster cell elongation on the shaded side than on the illuminated side. We now know that as light passes through a plant stem, it is diffracted and generates phototropin activation across the stem. Most activation occurs on the lit side, causing the plant hormone indole acetic acid (IAA) to accumulate on the shaded side. Stem cells elongate under influence of IAA.
Cryptochromes are another class of blue-light absorbing photoreceptors that also contain a flavin-based chromophore. Cryptochromes set the plants 24-hour activity cycle, also know as its circadian rhythem, using blue light cues. There is some evidence that cryptochromes work together with phototropins to mediate the phototropic response.
Plant Responses to Gravity
Whether or not they germinate in the light or in total darkness, shoots usually sprout up from the ground, and roots grow downward into the ground. A plant laid on its side in the dark will send shoots upward when given enough time. Gravitropism ensures that roots grow into the soil and that shoots grow toward sunlight. Growth of the shoot apical tip upward is called negative gravitropism, whereas growth of the roots downward is called positive gravitropism.
Amyloplasts (also known as statoliths) are specialized plastids that contain starch granules and settle downward in response to gravity. Amyloplasts are found in shoots and in specialized cells of the root cap. When a plant is tilted, the statoliths drop to the new bottom cell wall. A few hours later, the shoot or root will show growth in the new vertical direction.
The mechanism that mediates gravitropism is reasonably well understood. When amyloplasts settle to the bottom of the gravity-sensing cells in the root or shoot, they physically contact the endoplasmic reticulum (ER), causing the release of calcium ions from inside the ER. This calcium signaling in the cells causes polar transport of the plant hormone IAA to the bottom of the cell. In roots, a high concentration of IAA inhibits cell elongation. The effect slows growth on the lower side of the root, while cells develop normally on the upper side. IAA has the opposite effect in shoots, where a higher concentration at the lower side of the shoot stimulates cell expansion, causing the shoot to grow up. After the shoot or root begin to grow vertically, the amyloplasts return to their normal position. Other hypotheses—involving the entire cell in the gravitropism effect—have been proposed to explain why some mutants that lack amyloplasts may still exhibit a weak gravitropic response.
Plant Responses to Wind and Touch
The shoot of a pea plant winds around a trellis, while a tree grows on an angle in response to strong prevailing winds. These are examples of how plants respond to touch or wind.
The movement of a plant subjected to constant directional pressure is called thigmotropism, from the Greek words thigma meaning “touch,” and tropism implying “direction.” Tendrils are one example of this. The meristematic region of tendrils is very touch sensitive; light touch will evoke a quick coiling response. Cells in contact with a support surface contract, whereas cells on the opposite side of the support expand. Application of jasmonic acid is sufficient to trigger tendril coiling without a mechanical stimulus.
A thigmonastic response is a touch response independent of the direction of stimulus. In the Venus flytrap, two modified leaves are joined at a hinge and lined with thin fork-like tines along the outer edges. Tiny hairs are located inside the trap. When an insect brushes against these trigger hairs, touching two or more of them in succession, the leaves close quickly, trapping the prey. Glands on the leaf surface secrete enzymes that slowly digest the insect. The released nutrients are absorbed by the leaves, which reopen for the next meal.
Thigmomorphogenesis is a slow developmental change in the shape of a plant subjected to continuous mechanical stress. When trees bend in the wind, for example, growth is usually stunted and the trunk thickens. Strengthening tissue, especially xylem, is produced to add stiffness to resist the wind’s force. Researchers hypothesize that mechanical strain induces growth and differentiation to strengthen the tissues. Ethylene and jasmonate are likely involved in thigmomorphogenesis.
Defense Responses against Herbivores and Pathogens
Plants face two types of enemies: herbivores and pathogens. Herbivores both large and small use plants as food, and actively chew them. Pathogens are agents of disease. These infectious microorganisms, such as fungi, bacteria, and nematodes, live off of the plant and damage its tissues. Plants have developed a variety of strategies to discourage or kill attackers.
The first line of defense in plants is an intact and impenetrable barrier. Bark and the waxy cuticle can protect against predators. Other adaptations against herbivory include thorns, which are modified branches, and spines, which are modified leaves. They discourage animals by causing physical damage and inducing rashes and allergic reactions. A plant’s exterior protection can be compromised by mechanical damage, which may provide an entry point for pathogens. If the first line of defense is breached, the plant must resort to a different set of defense mechanisms, such as toxins and enzymes.
Secondary metabolites are compounds that are not directly derived from photosynthesis and are not necessary for respiration or plant growth and development. Many metabolites are toxic, and can even be lethal to animals that ingest them. Some metabolites are alkaloids, which discourage predators with noxious odors (such as the volatile oils of mint and sage) or repellent tastes (like the bitterness of quinine). Other alkaloids affect herbivores by causing either excessive stimulation (caffeine is one example) or the lethargy associated with opioids. Some compounds become toxic after ingestion; for instance, glycol cyanide in the cassava root releases cyanide only upon ingestion by the herbivore.
Mechanical wounding and predator attacks activate defense and protection mechanisms both in the damaged tissue and at sites farther from the injury location. Some defense reactions occur within minutes: others over several hours. The infected and surrounding cells may die, thereby stopping the spread of infection.
Long-distance signaling elicits a systemic response aimed at deterring the predator. As tissue is damaged, jasmonates may promote the synthesis of compounds that are toxic to predators. Jasmonates also elicit the synthesis of volatile compounds that attract parasitoids, which are insects that spend their developing stages in or on another insect, and eventually kill their host. The plant may activate abscission of injured tissue if it is damaged beyond repair.
Check Your Understanding
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