Root Modifications Storage Roots Roots frequently provide long term storage for carbohydrates that accumulate during

summer photosynthesis. In biennial species (which lives for 2 years; for example, beets, carrots, and celery) and many perennials (Datura, Phlox, many daisies), roots are the only permanent organs: In autumn, most of the stem dies back to a few nodes located below ground at the top of the root. Carbohydrates stored in the root are used to produce a new shoot in the spring, when photosynthesis is impossible owing to the lack of leaves. It seems less economical in these species to winterproof the shoot than to replace it using nutrients stored in the roots. Annual plants can survive without such storage capacity. Perennial plants with permanent shoots that do not die back also store significant amounts of nutrients within themselves during winter, but roots offer certain advantages. Being subterranean, roots are less available as food than are swollen, highly nutritious, easily visible stems. Roots also have a much more stable environment, subjected to less extreme changes in temperature and humidity, which may be important for survival of storage parenchyma cells.

Prop Roots The stem of a monocot can become wider, with more vascular bundles, if it can produce adventitious roots that extend to the soil. In some cases, roots are capable of extensive growth through the air: In many palms, the exposed roots can be 20 to 50 cm long. In the screwpine (Pandanus, a monocot, not a pine), they are often 3 or 4 meters long and the roots may grow through the air for months before reaching the ground corn and many other grasses. After prop roots do make contact with the soil, they transport additional nutrients and water to the stem. Just as importantly, they contract slightly and place some tension on the stem, thus acting as stabilizers, much like guy wires on tall television antennas. If the roots undergo secondary growth and become woody, they can be extremely strong supports, permitting a branch to extend even farther from the trunk without breaking or sagging. In banyan trees (eudicots of the genus Ficus), prop roots and branches can spread and produce massive tress many meters in diameter. Roots of certain tropical tress become tall, plate-like buttress roots; their upper side grows more rapidly than other parts of the root. Buttress roots brace the trunk against being blown over by wind. Mangroves also have prop roots, but these seem to be selectively advantageous for other reasons. The plants grow in intertidal marshes and are subjected to powerful water currents during storms and even normal tide changes; brace-like prop roots provide much more stability than a taproot system would. In addition the aerial

portion of the prop root is covered with numerous air chambers-lenticels-and its cortex is a wide aerenchyma. The subterranean portion of the root grows in a stagnant muck that has little or no oxygen; it is able to respire only because the aerenchyma permits rapid diffusion of oxygen from the aerial lenticels to the submerged root tissues. If mangrove roots were entirely subterranean, respiration would be extremely reduced. Aerial Roots of Orchids Many orchids are epiphytic, living attached to the branches of trees. Their roots spread along the surface of the bark and often dangle freely in the air. Although these plants live in rainforests, the orchids are actually adapted to drought conditions. In the few hours when rain does not fall, the air and bark become dry and could easily pull water out of the orchids roots if there were no waterconserving mechanism. The root epidermis, called a velamen in these orchids, is composed of several layers of large dead cells that are white in appearance. Apparently, the velamen acts as a waterproof barrier, not permitting water to leave the sides of the root. Contractile Roots In Oxalis, Gladiolus, Crinum and other plants (many with bulbs), roots undergo even more contraction than prop roots do. After extending through the soil and becoming firmly anchored, the uppermost portions slowly contract. Because the root is firmly fixed to the soil, the stem is pulled downward so that the base of the shoot is either kept at soil level or, in the case of bulbs, actually buried deeper. The contraction is caused by changes in the shape of cortex cells. They simultaneously shorten and expand radially, losing as much as one half to two thirds of their height. The vascular tissues buckle and become undulate but are able to continue conducting. Contractile roots may be more common than is generally appreciated. Many seeds germinate at or near the soil surface; root contraction may be the means by which the shoot becomes anchored in the soil. In the bulbs, corms, rhizomes, and other subterranean stems, contractile roots can be important in keeping the stems at the proper depth.

Storage Roots. Plants may develop specialized thickened roots as those found on carrot (Daucus, f), beet (Beta), sweet potato (Ipomea batatas), manroot (Ipomea pandurata), tuberous begonia (Begonia), and dahlia (Dahlia). These roots are an adaptation for food storage and are seen in many biennials and perennials. A biennial plant is one that produces leaves the first year and flowers and fruit the second year, then dies. A perennial plant lives for more than two years. Prop Roots. Some plants develop supporting roots as found in corn (Zea, h) and mangrove (Rhizophora). Such roots are especially effective in anchoring the shoot system in the soil, preventing them from capsizing in strong winds, heavy rains, and impact from predators and human activites. Contractile Roots. Contraction, or shortening of some roots, helps pull down and anchor the plant more firmly in the soil. Contractile roots are in herbaceous dicots and monocots, and occur in taproots, adventitious roots, and lateral roots, and on roots of underground storage stems such as bulbs and corms. (An herbaceous plant or herb is nonwoody and can die to the ground in freezing climates.) Nodulated tap roots These are roots modified for N2 fixation. These roots are observed in leguminous plants (pea, gram). These roots bear many small irregular swellings called root nodules. These nodules are living places for nitrogen fixing bacteria, Rhizobium leguminosarum. These bacteria fix and convert free atmospheric nitrogen into nitrates which are absorbed by roots. Pneumatophores or respiratory roots These are aerial (-ively geotropic) roots or aerophores produced in mangrove plants e.g. Sonneratia, Avicennia, Rhizophora. The underground roots of plants sent out aerial roots or Pneumatophores. These bear several pores or lenticels which help in gaseous exchange.

Underground Root Modifications

Taproot and adventitious roots may undergo certain modifications to perform the function of storage and vegetative propagation.

Tap Root Modifications (for storage of food)

In some plants the tap roots store reserve food for which they become swollen and assume different shapes. There are four types :

Fusiform root - In radish the modified tap root is thickened in the middle and tapers towards both ends.

e.g., Raphanus sativus (Radish)

Napiform root - In beetroot the modified tap root is greatly swollen at the base, but abruptly narrows into a tail-like portion.

e.g., Beta vulgaris (Beetroot)

Conical root - In carrot the modified tap root is conical. It is broad at the base and gradually tapers towards its apex.

e.g., Daucas carota (carrot)

fig. 27.5 - Tap Root Modifications

Adventitious Root Modifications

Tuberous or tubercular root, e.g., Ipomoea batotas (Sweet potato)

Here, the modified adventitious root is swollen without any definite shape. It is always produced singly and not in clusters.

Fasciculated roots e.g., Asparagus, Dahlia

In Dahlia and Asparagus a number of adventitious roots arise as a cluster, from almost the same level at the base of the stem.

Nodulose roots e.g., Curcuma amada, (mango ginger)

Here, the root becomes swollen at its tip.

Moniliform or beaded roots e.g., Momordica

Here, the roots show beadlike swellings at frequent intervals.

Annulated roots E.g., Ipecac

Here, the adventitious roots have a series of ring-like swellings on their surface.

fig. 27.6 - Modified Adventitious Roots (for storage of food)

Aerial Root Modifications

Aerial roots are adventitious roots which develop from the aerial parts of the plant to perform various functions.

A. For Mechanical Support

Prop roots e.g., Ficus benghalensis (Banyan) : In banyan the adventitious roots arise from the horizontal branches and grow vertically downwards. After reaching the soil they become thick and woody. Thus, they function as pillars giving mechanical support to the branches. Hence, they are also known as columnar roots.

fig. 27.7 - Modified Adventitious Roots (for mechanical support)

Stilt roots e.g. Pandanus : In Pandanus the adventitious roots arise from the lower part of the main stem and grow obliquely towards the soil. They serve to keep the plant erect by giving additional support. Climbing roots or clinging roots e.g., Piper betel, Pothos, etc : These roots arise from the nodes and help in attaching the climbing stem firmly to a support like a tree or a wall, by various mechanisms.

B. For Vital Functions

Haustoria or sucking roots e.g. Cuscuta : Parasites like Cuscuta develop a kind of root which penetrates into the tissue of the host plant and help to draw nutrients from the host by sucking it. The parasitic plants are not completely equipped to prepare their food. Hence, such plants have to depend on host plants for nutrients.

fig. 27.8 - Modified Adventitious Roots: Cuscuta (For vital functions)

Respiratory roots or breathing roots or pneumtophores.

e.g., Avicennia, Rhizophora.

fig. 27.9: Pneumatophores A number of plants growing in marshy water-logged soils which contain almost no air, develop some branches which grow vertically upwards into the air. These roots are called breathing roots or pneumatophores. Each such root is provided towards the upper end with numerous pores through which gases diffuse in and out.

Floating roots e.g., Jussiea: In Jussiea which is an aquatic plant, special spongy roots called floating or respiratory roots arise from the plant. They are adventitious and enclose a tissue called aerenchyma. These roots usually develop above the level of water and serve to store up air and help in buoyancy of the plant. In addition, the floating roots may also perform the respiratory function. Hence, they are also called respiratory roots. Epiphytic roots e.g., Vanda: In epiphytes like orchids which grow on other plants, special adventitious roots called epiphytic roots are produced. The outer region of the root is made up of a special tissue called the velamen. This tissue absorbs moisture from the air

and makes it available to the plant. These roots do not penetrate the host tissue like parasitic roots.

Summary of Root Modifications

Underground Stem Modifications

The underground stems, by being situated below the surface of the soil, protect themselves against unfavourable conditions of weather and the attack of animals, and serve as store houses for reserve food, and in vegetative propagation. Their stem nature can be distinguished by the presence of nodes and internodes, scale leaves at the nodes, axillary buds in axils of scale leaves and a terminal bud. Further, the anatomy of the underground stem resembles that of an aerial stem. The underground stems are of four types namely rhizome, tuber, bulb and corm.

Rhizome
A rhizome is a thick horizontally growing stem which usually stores food material. It has nodes and internodes, scale leaves, axillary buds, adventitous roots and a terminal bud. Scale leaves enclosing the axillary buds are seen arising from the nodal points of the stem. Some of the axillary buds develop into branches which

grow upwards into the air and then produce normal green foliage leaves. Usually the growing points of the rhizome continue to remain underground causing an elongation of the rhizome. Roots develop from the lower surface of the rhizome. Eg. Ginger, Turmeric.

fig. 27.17 Rhizome of Ginger

fig. 27.18 Root Stock (Vertical Rhizome) of Alocasia

Tuber
Tuber is a swollen end of an underground branch which arises from the axil of a lower leaf. These underground branches grow horizontally outwards in the soil. Each

tuber is irregular in shape due to the deposition of food materials (starch). On the surface of each tuber many leaf scars are seen. These leaf scars are the impressions of fallen scale leaves. Each such leaf scar encloses an axillary bud. A leaf scar with an axillary bud is called an eye. These eyes of potato are capable of producing new plants by vegetative propagation. E.g., Potato.

fig. 27.19 Tuber of Potato

Bulb
Here, the stem is reduced and represented by a short disc. The lower surface of the stem produces many adventitious roots. E.g., Onion, Garlic.
In bulbs of onion, garlic, etc. the inner leaves are fleshy while the outer ones are dry. This is called as tunicated bulb since the concentric leaf bases form a complete covering or tunic. The apical bud of the bulb produces the shoot. The axillary buds sometimes produce daughter bulbs, as in garlic.

fig. 27.20 Bulb of Onion

fig. 27.20 Bulb of garlic

Corm
A corm is a greatly swollen underground basal portion of an erect stem. The swelling is due to the storage of reserve food material. It bears scale leaves and axillary buds. At the end of the growing season, the aerial parts die. With the return of favorable conditions usually one axillary bud (rarely more than one) near the apex develops into a new shoot utilising the food reserve material in the old corm. The new plant produces a new corm at its base. The earlier corm shrivels off. E.g. Amorphophallus, Colocasia.

fig. 27.21 Corm of Colocasia

Subaerial Stem Modifications

In some plants, the sub-aerial stems are modified for the purpose of vegetative reproduction. They are of the following types :

Runner

The runner arises from the base of the stem as a lateral branch and runs along the surface of the soil. It develops distinct nodes and internodes. At each node, the runner produces roots below and leaves above. In this way many runners are often produced by the mother plant and they spread out on the ground on all sides. If any accidental injury results in the separation of a runner, the severed parts are capable of leading an independent existence. E.g., Oxalis, Fragaria, Centella astatica.

fig. 27.22 - Runners

Offset
An offset is a short thick runner like branch which produces a new plant at its tip. The offsets grow in all directions from the main stem of the parent plant. If any accidental injury results in the separation of these units, each is capable of leading an independent existence. E.g., Pistia, Eichhornia.

fig. 27.23 Offsets

Stolon

Here, lateral branches called stolons originate from the underground stem. The stolons grow horizontally outwards for a varying distance in the soil. Ultimately their end (terminal bud) emerges out of the ground and develops into a new plant. A runner, sucker or any basal branch which produces roots is called a stolon. E.g. Colocasia.

fig. 27.24 Stolon

Sucker
A lateral branch arising close to the ground level, traveling underground for some distance, turning up at its end and producing a new plant is a sucker. Eg. Chrysanthemum

fig. 27.25 Sucker

Modifications of Leaves

A normal leaf is thin, flat and green and performs the function of photosynthesis. In some plants certain special functions are performed by leaves, which become modified.

Leaf Tendrils
Tendrils are slender, spirally coiled springlike structures. They are highly sensitive to contact and when they come in contact with any support, tendrils coil around the support like the stem twiners.

In glory lily (Gloriosa superba) the leaf apex is modified into a tendril. In pea (Pisum sativum) the terminal leaflets of an unipinnately compound leaf are modified into tendrils. In Lathyrus or wild pea, the entire leaf is modified into a tendril. In Clematis and Smilax, the petiole and stipules respectively, are modified into tendrils.

Phyllodes
A phyllode is the petiole or rachis of a leaf which is modified into a green flat structure for the purpose of photosynthesis. In such a leaf the lamina is poorly developed. In Acacia melanoxylon, the petiole is flattened, green and becomes a phyllode. The leaflets and secondary rachii drop off.

In Parkinsonia aculeata, the secondary rachii are modified into phylodes which are photosynthetic. The primary rachis is modified into a spine.

Leaf Spines
In some plants, leaves or parts of leaves may be modified into spines.

In Opuntia (prickly pear) leaves are poorly developed and fall of very early, but the minute leaves of the axillary bud are modified into spines. In Argemone (prickly poppy), the leaf margin is modified into small spines. In Zizyphus the stipules are modified into spines. The spines act as defensive structures.

fig. 27.38 - Leaf Spines

Scale Leaves
In many desert plants, the leaves are highly reduced and appear as scales. The scale leaves are thin, membranous, dry, stalkless and brownish or colourless. In plants where the leaves are reduced to scales in order to minimise transpiration, the function of photosynthesis is relegated to the stems (cladodes).