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Tuber

Ulluku (Ullucus tuberosus) Tubers are enlarged structures in some species used as storage organs for nutrients. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growing season, and as a means of .[1] Stem tubers form from thickened (underground stems) or (horizontal connections between organisms). Common plant species with stem tubers include and . Some sources also treat modified lateral ( tubers) under the definition; these are encountered in , , and dahlia. Terminology [edit] The term originates from Latin , meaning "lump, bump, swelling".[2] Some sources define the term "tuber" to mean only structures derived from stems;[3] others use the term for structures derived from stems or roots.[4] Stem tubers[edit] A stem tuber forms from thickened rhizomes or stolons. The top sides of the tuber produce shoots that grow into typical stems and leaves and the under sides produce roots. They tend to form at the sides of the parent plant and are most often located near the soil surface. The tuber is normally a short-lived storage and regenerative organ developing from a shoot that branches off a mature plant. The offsprings or new tubers are attached to a parent tuber or form at the end of a hypogeogenous (initiated below ground) . In the autumn the plant dies, except for the new offspring stem tubers which have one dominant , which in spring regrows a new shoot producing stems and leaves, in summer the tubers decay and new tubers begin to grow. Some also form smaller tubers and/or tubercules which act like , producing small plants that resemble (in morphology and size) seedlings. Some stem tubers are long-lived, such as those of tuberous begonia, but many plants have tubers that survive only until the plants have fully leafed out, at which point the tuber is reduced to a shriveled-up husk.

Flowers and tuber of Anredera cordifolia Stem tubers generally start off as enlargements of the hypocotyl section of a seedling but also sometimes include the first node or two of the epicotyl and the upper section of the root. The stem tuber has a vertical orientation with one or a few vegetative on the top and fibrous roots produced on the bottom from a basal section, typically the stem tuber has an oblong rounded shape. Tuberous begonia, yams,[5][6] and are commonly grown stem tubers. Mignonette vine (Anredera cordifolia) produces aerial stem tubers on 12-to-25-foot-tall (3.7 to 7.6 m) vines, the tubers fall to the ground and grow. Plectranthus esculentus of the mint family Lamiaceae, produces tuberous under ground organs from the base of the stem, weighing up to 1.8 kg per tuber, forming from axillary buds producing short stolons that grow into tubers.[7] Potatoes[edit] Main article: Potato

Potato plant with revealed tubers Potatoes are stem tubers. Enlarged stolons thicken to develop into storage organs.[8][9][10][Links have expired] The tuber has all the parts of a normal stem, including nodes and internodes. The nodes are the eyes and each has a leaf scar. The nodes or eyes are arranged around the tuber in a spiral fashion beginning on the end opposite the attachment point to the . The terminal bud is produced at the farthest point away from the stolon attachment and tubers thus show the same apical dominance as a normal stem. Internally, a tuber is filled with stored in enlarged parenchyma like cells. The inside of a tuber has the typical cell structures of any stem, including a pith, vascular zones, and a cortex. The tuber is produced in one growing season and used to perennate the plant and as a means of propagation. When fall comes, the above-ground structure of the plant dies, but the tubers survive over winter underground until spring, when they regenerate new shoots that use the stored in the tuber to grow. As the main shoot develops from the tuber, the base of the shoot close to the tuber produces adventitious roots and lateral buds on the shoot. The shoot also produces stolons that are long etiolated stems. The stolon elongates during long days with the presence of high auxins levels that prevent root growth off of the stolon. Before new tuber formation begins, the stolon must be a certain age. The enzyme lipoxygenase makes a hormone, jasmonic acid, which is involved in the control of potato tuber development. The stolons are easily recognized when potato plants are grown from seeds. As the plants grow, stolons are produced around the soil surface from the nodes. The tubers form close to the soil surface and sometimes even on top of the ground. When potatoes are cultivated, the tubers are cut into pieces and planted much deeper into the soil. Planting the pieces deeper creates more area for the plants to generate the tubers and their size increases. The pieces sprout shoots that grow to the surface. These shoots are rhizome-like and generate short stolons from the nodes while in the ground. When the shoots reach the soil surface, they produce roots and shoots that grow into the green plant. Root tubers[edit]

Freshly dug sweet potato plants with tubers.

A tuberous root or storage root, is a modified lateral root, enlarged to function as a . The enlarged area of the root-tuber, or storage root, can be produced at the end or middle of a root or involve the entire root. It is thus different in origin but similar in function and appearance to a stem tuber. Examples of plants with notable tuberous roots include the sweet potato, cassava, and dahlia. Root tubers are perennating organs, thickened roots that store nutrients over periods when the plant cannot actively grow, thus permitting survival from one year to the next. The massive enlargement of secondary roots typically represented by sweet potato (Ipomoea batatas), have the internal and external cell and tissue structures of a normal root, they produce adventitious roots and stems which again produce adventitious roots.[11]

In root-tubers, there are no nodes and internodes or reduced leaves. Root tubers have one end called the proximal end, which is the end that was attached to the old plant; this end has crown tissue that produces buds which grow into new stems and foliage.[12] The other end of the root tuber is called the distal end, and it normally produces unmodified roots. In stem tubers the order is reversed, with the distal end producing stems. Tuberous roots are biennial in duration: the first year the plant produces root-tubers, and at the end of the growing season, the plant shoots often die, leaving the newly generated tubers. The next growing season, the root-tubers produce new shoots. As the shoots of the new plant grow, the stored reserves of the root-tuber are consumed in the production of new roots, stems, and reproductive organs; any remaining root tissue dies concurrently to the plant's regeneration of next generation of root-tubers.

Hemerocallis fulva plus a number of Daylily hybrids have large root-tubers, H. fulva spreads by underground stolons[13] that end with a new fan that grows roots that produce thick root tubers and then send out more stolons.[14][15]

Root tubers, along with other storage tissues that plants produce, are consumed by animals as a rich source of nutrients. The root-tubers of Arrowhead plants of the genus Sagittaria are eaten by ducks.[16]

Plants with root tubers are propagated in late summer to late winter by digging up the tubers and separating them, making sure that each piece has some crown tissue for replanting.