Cactus Pdf

[Yi Pressimone]

Acactus (pl.: cacti, cactuses, or less commonly, cactus)[3] is a member of the plant family Cactaceae (/kˈkteɪsiaɪ, -siːiː/),[a] a family comprising about 127 genera with some 1,750 known species of the order Caryophyllales.[4] The word cactus derives, through Latin, from the Ancient Greek word κάκτος (kktos), a name originally used by Theophrastus for a spiny plant whose identity is now not certain.[5] Cacti occur in a wide range of shapes and sizes. They are native to the Americas, ranging from Patagonia in the south to parts of western Canada in the north, with the exception of Rhipsalis baccifera, which is also found in Africa and Sri Lanka. Cacti are adapted to live in very dry environments, including the Atacama Desert, one of the driest places on Earth. Because of this, cacti show many adaptations to conserve water. For example, almost all cacti are succulents, meaning they have thickened, fleshy parts adapted to store water. Unlike many other succulents, the stem is the only part of most cacti where this vital process takes place. Most species of cacti have lost true leaves, retaining only spines, which are highly modified leaves. As well as defending against herbivores, spines help prevent water loss by reducing air flow close to the cactus and providing some shade. In the absence of true leaves, cacti's enlarged stems carry out photosynthesis.

Cactus spines are produced from specialized structures called areoles, a kind of highly reduced branch. Areoles are an identifying feature of cacti. As well as spines, areoles give rise to flowers, which are usually tubular and multipetaled. Many cacti have short growing seasons and long dormancies and are able to react quickly to any rainfall, helped by an extensive but relatively shallow root system that quickly absorbs any water reaching the ground surface. Cactus stems are often ribbed or fluted with a number of ribs which corresponds to a number in the Fibonacci numbers (2, 3, 5, 8, 13, 21, 34 etc.). This allows them to expand and contract easily for quick water absorption after rain, followed by retention over long drought periods. Like other succulent plants, most cacti employ a special mechanism called "crassulacean acid metabolism" (CAM) as part of photosynthesis. Transpiration, during which carbon dioxide enters the plant and water escapes, does not take place during the day at the same time as photosynthesis, but instead occurs at night. The plant stores the carbon dioxide it takes in as malic acid, retaining it until daylight returns, and only then using it in photosynthesis. Because transpiration takes place during the cooler, more humid night hours, water loss is significantly reduced.

Many smaller cacti have globe-shaped stems, combining the highest possible volume for water storage with the lowest possible surface area for water loss from transpiration. The tallest[b] free-standing cactus is Pachycereus pringlei, with a maximum recorded height of 19.2 m (63 ft),[7] and the smallest is Blossfeldia liliputiana, only about 1 cm (0.4 in) in diameter at maturity.[8] A fully grown saguaro (Carnegiea gigantea) is said to be able to absorb as much as 200 U.S. gallons (760 L; 170 imp gal) of water during a rainstorm.[9] A few species differ significantly in appearance from most of the family. At least superficially, plants of the genera Leuenbergeria, Rhodocactus and Pereskia resemble other trees and shrubs growing around them. They have persistent leaves, and when older, bark-covered stems. Their areoles identify them as cacti, and in spite of their appearance, they, too, have many adaptations for water conservation. Leuenbergeria is considered close to the ancestral species from which all cacti evolved. In tropical regions, other cacti grow as forest climbers and epiphytes (plants that grow on trees). Their stems are typically flattened, almost leaf-like in appearance, with fewer or even no spines, such as the well-known Christmas cactus or Thanksgiving cactus (in the genus Schlumbergera).

Cacti have a variety of uses: many species are used as ornamental plants, others are grown for fodder or forage, and others for food (particularly their fruit). Cochineal is the product of an insect that lives on some cacti.

The remaining cacti fall into only two groups: three tree-like genera, Leuenbergeria, Pereskia and Rhodocactus (all formerly placed in Pereskia), and the much smaller Maihuenia. These two groups are rather different from other cacti,[10] which means any description of cacti as a whole must frequently make exceptions for them. Species of the first three genera superficially resemble other tropical forest trees. When mature, they have woody stems that may be covered with bark and long-lasting leaves that provide the main means of photosynthesis. Their flowers may have superior ovaries (i.e., above the points of attachment of the sepals and petals) and areoles that produce further leaves. The two species of Maihuenia have succulent but non-photosynthetic stems and prominent succulent leaves.[10]

Cacti can be tree-like (arborescent), meaning they typically have a single more-or-less woody trunk topped by several to many branches. In the genera Leuenbergeria, Pereskia and Rhodocactus, the branches are covered with leaves, so the species of these genera may not be recognized as cacti. In most other cacti, the branches are more typically cactus-like, bare of leaves and bark and covered with spines, as in Pachycereus pringlei or the larger opuntias. Some cacti may become tree-sized but without branches, such as larger specimens of Echinocactus platyacanthus. Cacti may also be described as shrubby, with several stems coming from the ground or from branches very low down, such as in Stenocereus thurberi.[11]

Smaller cacti may be described as columnar. They consist of erect, cylinder-shaped stems, which may or may not branch, without a very clear division into trunk and branches. The boundary between columnar forms and tree-like or shrubby forms is difficult to define. Smaller and younger specimens of Cephalocereus senilis, for example, are columnar, whereas older and larger specimens may become tree-like. In some cases, the "columns" may be horizontal rather than vertical. Thus, Stenocereus eruca can be described as columnar even though it has stems growing along the ground, rooting at intervals.[11]

Cacti whose stems are even smaller may be described as globular (or globose). They consist of shorter, more ball-shaped stems than columnar cacti. Globular cacti may be solitary, such as Ferocactus latispinus, or their stems may form clusters that can create large mounds. All or some stems in a cluster may share a common root.[11]

Other cacti have a quite different appearance. In tropical regions, some grow as forest climbers and epiphytes. Their stems are typically flattened and almost leaf-like in appearance, with few or even no spines. Climbing cacti can be very large; a specimen of Hylocereus was reported as 100 meters (330 ft) long from root to the most distant stem. Epiphytic cacti, such as species of Rhipsalis or Schlumbergera, often hang downwards, forming dense clumps where they grow in trees high above the ground.[11]

The leafless, spiny stem is the characteristic feature of the majority of cacti (all belonging to the largest subfamily, the Cactoideae). The stem is typically succulent, meaning it is adapted to store water. The surface of the stem may be smooth (as in some species of Opuntia) or covered with protuberances of various kinds, which are usually called tubercles. These vary from small "bumps" to prominent, nipple-like shapes in the genus Mammillaria and outgrowths almost like leaves in Ariocarpus species. The stem may also be ribbed or fluted in shape. The prominence of these ribs depends on how much water the stem is storing: when full (up to 90% of the mass of a cactus may be water), the ribs may be almost invisible on the swollen stem, whereas when the cactus is short of water and the stems shrink, the ribs may be very visible.[11]

The stems of most cacti are some shade of green, often bluish or brownish green. Such stems contain chlorophyll and are able to carry out photosynthesis; they also have stomata (small structures that can open and close to allow passage of gases). Cactus stems are often visibly waxy.[11]

Areoles are structures unique to cacti. Although variable, they typically appear as woolly or hairy areas on the stems from which spines emerge. Flowers are also produced from areoles. In the genus Leuenbergeria, believed similar to the ancestor of all cacti, the areoles occur in the axils of leaves (i.e. in the angle between the leaf stalk and the stem).[12] In leafless cacti, areoles are often borne on raised areas on the stem where leaf bases would have been.

Areoles are highly specialized and very condensed shoots or branches. In a normal shoot, nodes bearing leaves or flowers would be separated by lengths of stem (internodes). In an areole, the nodes are so close together, they form a single structure. The areole may be circular, elongated into an oval shape, or even separated into two parts; the two parts may be visibly connected in some way (e.g. by a groove in the stem) or appear entirely separate (a dimorphic areole). The part nearer the top of the stem then produces flowers, the other part spines. Areoles often have multicellular hairs (trichomes) that give the areole a hairy or woolly appearance, sometimes of a distinct color such as yellow or brown.[11]

In most cacti, the areoles produce new spines or flowers only for a few years and then become inactive. This results in a relatively fixed number of spines, with flowers being produced only from the ends of stems, which are still growing and forming new areoles. In Pereskia, a genus close to the ancestor of cacti, areoles remain active for much longer; this is also the case in Opuntia and Neoraimondia.[11]

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