Plants play a key role in all of the biogeochemical cycles. Take the water cycle for example. Most water in the soil will be taken up by plant roots. The plant will use some of this water for its own metabolism, and some of that will find its way into animals that eat the plants, but much of it will be lost back to the atmosphere through a process known as evapotranspiration. Water enters the vascular system of the plant through the roots and evaporates, or transpires, through the stomata of the leaves. In the carbon cycle, plants are incredibly impactful. Carbon compounds contain energy, and many of these compounds from plants and algae have remained stored as fossilized carbon, which humans use as fuel. In the nitrogen and phosphorus cycles, plants take up nutrients from their environment. When they die and decompose, nutrients are released back into the environment. In this chapter, we will take a deep dive into the ways plants grow and interact with the nutrients in their environment.

Plant Root and Shoot Systems

In plants, just as in animals, similar cells working together form a tissue. When different types of tissues work together to perform a unique function, they form an organ; organs working together form organ systems. Vascular plants have two distinct organ systems: a shoot system, and a root system. The shoot system consists of two portions: the vegetative (non-reproductive) parts of the plant, such as the leaves and the stems, and the reproductive parts of the plant, which include flowers and fruits. The shoot system generally grows above ground, where it absorbs the light needed for photosynthesis. The root system, which supports the plants and absorbs water and minerals, is usually underground. Figure 27.1 shows the organ systems of a typical plant.

Shoot System

Stems

Stems may range in length from a few millimeters to hundreds of meters, and also vary in diameter, depending on the plant type. Stems are usually above ground, although the stems of some plants, such as the potato, also grow underground. Stems may be herbaceous (soft) or woody in nature. Their main function is to provide support to the plant, holding leaves, flowers and buds; in some cases, stems also store food for the plant. A stem may be unbranched, like that of a palm tree, or it may be highly branched, like that of a magnolia tree. The stem of the plant connects the roots to the leaves, helping to transport absorbed water and minerals to different parts of the plant. It also helps to transport the products of photosynthesis, namely sugars, from the leaves to the rest of the plant.

Plant stems, whether above or below ground, are characterized by the presence of nodes and internodes (Figure 27.2). Nodes are points of attachment for leaves, aerial roots, and flowers. The stem region between two nodes is called an internode. The stalk that extends from the stem to the base of the leaf is the petiole. An axillary bud is usually found in the axil—the area between the base of a leaf and the stem—where it can give rise to a branch or a flower. The apex (tip) of the shoot contains the apical meristem within the apical bud.

Leaves

Leaves are the main sites for photosynthesis: the process by which plants synthesize food. Most leaves are usually green, due to the presence of chlorophyll in the leaf cells. However, some leaves may have different colors, caused by other plant pigments that mask the green chlorophyll.

The epidermis of a leaf contains openings known as stomata, through which the exchange of gases takes place (Figure 27.3). Two cells, known as guard cells, surround each leaf stoma, controlling its opening and closing and thus regulating the uptake of carbon dioxide and the release of oxygen and water vapor. The thickness, shape, and size of leaves are adapted to the environment. Usually, the leaves of plants growing in tropical rainforests have larger surface areas than those of plants growing in deserts or very cold conditions, which are likely to have a smaller surface area to minimize water loss.

The epidermis is usually one cell layer thick; however, in plants that grow in very hot or very cold conditions, the epidermis may be several layers thick to protect against excessive water loss from transpiration. A waxy layer known as the cuticle covers the leaves of all plant species. The cuticle reduces the rate of water loss from the leaf surface.

Some leaves may have small hairs called trichomes on the leaf surface. Trichomes help to deter herbivory by restricting insect movements, or by storing toxic or bad-tasting compounds; they can also reduce the rate of water loss via transpiration by blocking air flow across the leaf surface.

The Root System

Root systems are mainly of two types: tap or fibrous (Figure 27.4). A tap root system has a main root that grows down vertically, and from which many smaller lateral roots arise. Dandelions are a good example; their tap roots usually break off when trying to pull these weeds, and they can regrow another shoot from the remaining root. A tap root system penetrates deep into the soil. In contrast, a fibrous root system is located closer to the soil surface, and forms a dense network of roots that also helps prevent soil erosion (lawn grasses are a good example, as are wheat, rice, and corn). Some plants have a combination of tap roots and fibrous roots. Plants that grow in dry areas often have deep root systems, whereas plants growing in areas with abundant water are likely to have shallower root systems. Root structures may be modified for specific purposes. For example, some roots are bulbous and store starch. Aerial roots and prop roots are two forms of aboveground roots that provide additional support to anchor the plant. Tap roots, such as carrots, turnips, and beets, are examples of roots that are modified for food storage.

Root hairs are extensions of root epidermal cells, increase the surface area of the root, greatly contributing to the absorption of water and minerals.

A layer of cells known as the endodermis separates the stele (vascular tissue) from the ground tissue in the outer portion of the root. The endodermis is exclusive to roots, and serves as a checkpoint for materials entering the root’s vascular system. A waxy substance called suberin is present on the walls of the endodermal cells. This waxy region, known as the Casparian strip, forces water and solutes to cross the plasma membranes of endodermal cells instead of slipping between the cells. This ensures that only materials required by the root pass through the endodermis, while toxic substances and pathogens are generally excluded.

Plant Growth

Growth in plants occurs as the stems and roots lengthen. Some plants, especially those that are woody, also increase in thickness during their life span. The increase in length of the shoot and the root is referred to as primary growth, and is the result of cell division in the shoot apical meristem. Secondary growth is characterized by an increase in thickness or girth of the plant, and is caused by cell division in the lateral meristem. Herbaceous plants mostly undergo primary growth, with hardly any secondary growth or increase in thickness. Secondary growth or “wood” is noticeable in woody plants; it occurs in some dicots, but occurs very rarely in monocots.

Meristems are plant regions of continuous cell division and growth. Meristems produce cells that quickly differentiate, or specialize, and become permanent tissue. Meristematic tissues consist of three types, based on their location in the plant. Apical meristems are located at the tips of stems and roots, which enable a plant to extend in length (primary growth). Lateral meristems facilitate growth in thickness or girth in a maturing plant (secondary growth). Intercalary meristems occur only in monocots, at the bases of leaf blades and at nodes (the areas where leaves attach to a stem). This tissue enables the monocot leaf blade to increase in length from the leaf base; for example, it allows lawn grass leaves to elongate even after repeated mowing.

Some plant parts, such as stems and roots, continue to grow throughout a plant’s life: a phenomenon called indeterminate growth. Other plant parts, such as leaves and flowers, exhibit determinate growth, which ceases when a plant part reaches a particular size.

Primary Growth

Most primary growth occurs at the apices, or tips, of stems and roots. Primary growth is a result of rapidly dividing cells in the apical meristems at the shoot tip and root tip. Subsequent cell elongation also contributes to primary growth. The growth of shoots and roots during primary growth enables plants to continuously seek water (roots) or sunlight (shoots).

The influence of the apical bud on overall plant growth is known as apical dominance, which diminishes the growth of axillary buds that form along the sides of branches and stems. Most coniferous trees exhibit strong apical dominance, thus producing the typical conical Christmas tree shape. If the apical bud is removed, then the axillary buds will start forming lateral branches. Gardeners make use of this fact when they prune plants by cutting off the tops of branches, thus encouraging the axillary buds to grow out, giving the plant a bushy shape.

Shoots and roots of plants increase in length through rapid cell division in a tissue called the apical meristem, which is a small mitotically active zone of cells found at the shoot tip or root tip. The apical meristem is made of undifferentiated cells that continue to proliferate throughout the life of the plant. Meristematic cells give rise to all the specialized tissues of the organism. Elongation of the shoots and roots allows a plant to access additional space and resources: light in the case of the shoot, and water and minerals in the case of roots.

Secondary Growth

The increase in stem thickness that results from secondary growth is due to the activity of the lateral meristems, which are lacking in herbaceous plants. Lateral meristems include the vascular cambium and, in woody plants, the cork cambium. The vascular cambium is located just outside the primary xylem and to the interior of the primary phloem. The cells of the vascular cambium divide and form secondary xylem (tracheids and vessel elements) to the inside, and secondary phloem (sieve elements and companion cells) to the outside. The thickening of the stem that occurs in secondary growth is due to the formation of secondary phloem and secondary xylem by the vascular cambium, plus the action of cork cambium, which forms the tough outermost layer of the stem. The cells of the secondary xylem contain lignin, which provides hardiness and strength.

In woody plants, cork cambium is the outermost lateral meristem. It produces cork cells (bark) containing a waxy substance known as suberin that can repel water. The bark protects the plant against physical damage and helps reduce water loss. In some plants, the bark has many openings which allow the interior cells to exchange gases with the outside atmosphere. This supplies oxygen to the living and metabolically active cells of the cortex, xylem, and phloem.

The activity of the vascular cambium gives rise to annual growth rings. During the spring growing season, cells of the secondary xylem have a large internal diameter and their primary cell walls are not extensively thickened. This is known as early wood, or spring wood. During the fall season, the secondary xylem develops thickened cell walls, forming late wood, or autumn wood, which is denser than early wood. This alternation of early and late wood is due largely to a seasonal decrease in the number of vessel elements and a seasonal increase in the number of tracheids. It results in the formation of an annual ring, which can be seen as a circular ring in the cross section of the stem (Figure 27.5). An examination of the number of annual rings and their nature (such as their size and cell wall thickness) can reveal the age of the tree and the prevailing climatic conditions during each season.

Phototropism

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.

A plant’s sensory response to external stimuli relies on chemical messengers (hormones). Plant hormones affect all aspects of plant life, from flowering to fruit setting and maturation, and from phototropism to leaf fall. Potentially every cell in a plant can produce plant hormones. They can act in their cell of origin or be transported to other portions of the plant body, with many plant responses involving the synergistic or antagonistic interaction of two or more hormones. In contrast, animal hormones are produced in specific glands and transported to a distant site for action, and they act alone.

Auxin is a plant hormone related to growth, including phototropism (in fact, the term auxin is derived from the Greek word auxein, which means “to grow”). Auxins also control the differentiation of meristem into vascular tissue, and promote leaf development and arrangement. Apical dominance—the inhibition of lateral bud formation—is triggered by auxins produced in the apical meristem. Flowering, fruit setting and ripening, and inhibition of abscission (leaf falling) are other plant responses under the direct or indirect control of auxins. Auxins also act as a relay for the effects of the blue light and red/far-red responses.

Commercial use of auxins is widespread in plant nurseries and for crop production. Auxins are used as a rooting hormone to promote growth of adventitious roots on cuttings and detached leaves. Applying synthetic auxins to tomato plants in greenhouses promotes normal fruit development. Outdoor application of auxin promotes synchronization of fruit setting and dropping to coordinate the harvesting season. Fruits such as seedless cucumbers can be induced to set fruit by treating unfertilized plant flowers with auxins.

Plant Tissues

Plant cells grow from meristems and differentiate into three main types of tissues: dermal, vascular, and ground tissue. Dermal tissue covers and protects the plant (like skin on animals). Vascular tissue transports water, minerals, and sugars to different parts of the plant. Ground tissue serves as a site for photosynthesis, provides a supporting matrix for the vascular tissue, and helps to store water and sugars.

These tissues are formed from three types of cells: parenchyma, collenchyma, and sclerenchyma cells.

Parenchyma cells are the most common plant cells. They are found in the stem, the root, the inside of the leaf, and the pulp of the fruit. Parenchyma cells are responsible for metabolic functions, such as photosynthesis, and they help repair and heal wounds. Some parenchyma cells also store starch.

Collenchyma cells are elongated cells with unevenly thickened walls. They provide structural support, mainly to the stem and leaves. These cells are alive at maturity and are usually found below the epidermis. The “strings” of a celery stalk are an example of collenchyma cells.

Sclerenchyma cells also provide support to the plant, but unlike collenchyma cells, many of them are dead at maturity. There are two types of sclerenchyma cells: fibers and sclereids. Both types have secondary cell walls that are thickened with deposits of lignin, an organic compound that is a key component of wood. Fibers are long, slender cells; sclereids are smaller-sized. Sclereids give pears their gritty texture. Humans use sclerenchyma fibers to make linen and rope.

Vascular tissues

Vascular tissue is made of two specialized conducting tissues: xylem and phloem. Xylem tissue transports water and nutrients from the roots to different parts of the plant. Xylem includes three different cell types: vessel elements, tracheids (both of which conduct water), and xylem parenchyma. Phloem tissue transports organic compounds from the site of photosynthesis to other parts of the plant. Phloem consists of four different cell types: sieve cells (which conduct photosynthates), companion cells, phloem parenchyma, and phloem fibers. Xylem cells are dead, but phloem cells are alive at maturity. The xylem and phloem always lie adjacent to each other (Figure 27.6). In stems, the xylem and the phloem form a structure called a vascular bundle; in roots, this is termed the vascular stele or vascular cylinder.

The xylem and phloem that make up the vascular tissue of the stem are arranged in distinct strands called vascular bundles, which run up and down the length of the stem. When the stem is viewed in cross section, the vascular bundles of dicot stems are arranged in a ring. In plants with stems that live for more than one year, the individual bundles grow together and produce the characteristic growth rings. In monocot stems, the vascular bundles are randomly scattered throughout the ground tissue.

Xylem tissue has three types of cells: xylem parenchyma, tracheids, and vessel elements. The latter two types conduct water and are dead at maturity. Tracheids are xylem cells with thick secondary cell walls that are lignified. Water moves from one tracheid to another through regions on the side walls known as pits, where secondary walls are absent. Vessel elements are xylem cells with thinner walls; they are shorter than tracheids. Each vessel element is connected to the next by means of a perforation plate at the end walls of the element. Water moves through the perforation plates to travel up the plant.

Phloem tissue is composed of sieve-tube cells, companion cells, phloem parenchyma, and phloem fibers. A series of sieve-tube cells (also called sieve-tube elements) are arranged end to end to make up a long sieve tube, which transports organic substances such as sugars and amino acids. The sugars flow from one sieve-tube cell to the next through perforated sieve plates, which are found at the end junctions between two cells. Although still alive at maturity, the nucleus and other cell components of the sieve-tube cells have disintegrated. Companion cells are found alongside the sieve-tube cells, providing them with metabolic support. The companion cells contain more ribosomes and mitochondria than the sieve-tube cells, which lack some cellular organelles.

Movement of Water and Minerals in the Xylem

Plants are phenomenal hydraulic engineers. Using only the basic laws of physics and the simple manipulation of potential energy, plants can move water to the top of a 116-meter-tall tree (Figure 27.7a). Plants can also use hydraulics to generate enough force to split rocks and buckle sidewalks (Figure 27.7b). Plants achieve this because of water potential.

Solutes, pressure, and gravity are all important for the transport of water in plants. Transpiration is the loss of water from the plant through evaporation at the leaf surface. It is the main driver of water movement in the xylem (Figure 27.8). Transpiration is caused by the evaporation of water at the leaf–atmosphere interface. Water from the roots is pulled up by this tension. At night, when stomata shut and transpiration stops, the water is held in the stem and leaf by the adhesion of water to the cell walls of the xylem vessels and tracheids, and the cohesion of water molecules to each other. This is called the cohesion–tension theory of sap ascent. Transpiration—the loss of water vapor to the atmosphere through stomata—is a passive process, meaning that metabolic energy in the form of ATP is not required for water movement. The energy driving transpiration is the difference in energy between the water in the soil and the water in the atmosphere. However, transpiration is tightly controlled.

Transportation of Photosynthates in the Phloem

Plants need an energy source to grow. In seeds and bulbs, food is stored in polymers (such as starch) that are converted by metabolic processes into sucrose for newly developing plants. Once green shoots and leaves are growing, plants are able to produce their own food by photosynthesizing. The products of photosynthesis are called photosynthates, which are usually in the form of simple sugars such as sucrose.

Structures that produce photosynthates for the growing plant are referred to as sources. Sugars produced in sources, such as leaves, need to be delivered to growing parts of the plant via the phloem in a process called translocation. The points of sugar delivery, such as roots, young shoots, and developing seeds, are called sinks. Seeds, tubers, and bulbs can be either a source or a sink, depending on the plant’s stage of development and the season.

The products from the source are usually translocated to the nearest sink through the phloem. For example, the highest leaves will send photosynthates upward to the growing shoot tip, whereas lower leaves will direct photosynthates downward to the roots. Intermediate leaves will send products in both directions, unlike the flow in the xylem, which is always unidirectional (soil to leaf to atmosphere). The pattern of photosynthate flow changes as the plant grows and develops. Photosynthates are directed primarily to the roots early on, to shoots and leaves during vegetative growth, and to seeds and fruits during reproductive development. They are also directed to tubers for storage.

Photosynthates, such as sucrose, are produced in the mesophyll cells of photosynthesizing leaves. From there they are translocated through the phloem to where they are used or stored. Mesophyll cells are connected by cytoplasmic channels. Photosynthates move through these channels to reach phloem sieve-tube elements (STEs) in the vascular bundles. From the mesophyll cells, the photosynthates are loaded into the phloem STEs. The sucrose is actively transported against its concentration gradient (a process requiring ATP) into the phloem cells.

Phloem STEs have reduced cytoplasmic contents, and are connected by a sieve plate with pores that allow for pressure-driven bulk flow, or translocation, of phloem sap. Companion cells are associated with STEs. They assist with metabolic activities and produce energy for the STEs (27.9).

Once in the phloem, the photosynthates are translocated to the closest sink. Phloem sap is an aqueous solution that contains up to 30% sugar, minerals, amino acids, and plant growth regulators. Sucrose concentration in the sink cells is lower than in the phloem STEs because the sink sucrose has been metabolized for growth, or converted to starch for storage or other polymers, such as cellulose, for structural integrity. Unloading at the sink end of the phloem tube occurs by either diffusion or active transport of sucrose molecules from an area of high concentration to one of low concentration. Water diffuses from the phloem by osmosis and is then transpired or recycled via the xylem back into the phloem sap.

References

Adapted from:

Clark, M.A., Douglas, M., and Choi, J. (2018). Biology 2e. OpenStax. Retrieved from https://openstax.org/books/biology-2e/pages/1-introduction