Transgenic Plants Genetically modified crops (GM crops, or biotech crops) are plants, the DNA of which has

been modified using genetic engineering techniques, which are then used in agriculture. Plants are also transgenically modified in scientific research; see genetically modified organism for discussion. Genetic engineering techniques are much more precise than mutagenesis (mutation breeding) where an organism is exposed to radiation or chemicals to create a non-specific but stable change. Other techniques by which humans modify food organisms include selective breeding; plant breeding, and animal breeding, and somaclonal variation. In most cases the aim is to introduce a new trait to the plant which does not occur naturally in this species. Examples include resistance to certain pests, diseases or environmental conditions, or the production of a certain nutrient or pharmaceutical agent. Critics have objected to GM crops per se on several grounds, including ecological concerns, and economic concerns raised by the fact these organisms are subject to intellectual property law. GM crops also are involved in controversies over GM food with respect to whether food produced from GM crops is safe and whether GM crops are needed to address the world's food needs. See the genetically modified food controversies article for discussion of issues about GM crops and GM food. This article covers GM crops and their use in agriculture. There are separate articles on other aspects of genetic engineering. The genetic engineering article focuses on history and methods of genetic engineering, and on applications of genetic engineering and of GMOs. The article on GMOs focuses on what organisms have been genetically engineered and for what purposes. The two articles cover much of the same ground but with different organizations (sorted by application in the genetic engineering article; sorted by organism in the GMO article). There are separate articles on genetically modified food itself, regulation, and controversies. GM crops grown today, or under experimental development, have been modified with traits intended to provide benefit to farmers, consumers, or industry. These traits include improved shelf life,disease resistance, stress resistance, herbicide resistance, pest resistance, production of useful goods such as biofuel or drugs, and ability to absorb toxins, for use in bioremediation of pollution. Due to high regulatory and research costs, the majority of genetically modified crops in agriculture consist of commodity crops, [46][47] such as soybean, maize, cotton and rapeseed. Recently, some research and development has been targeted to enhancement of crops that are locally important in developing countries, such as insect[48] [49] resistant cowpea for Africa and insect-resistantbrinjal (eggplant) for India.
[1]

Improved shelf life

The first genetically modified crop approved for sale in the U.S. was the FlavrSavr tomato, which had a [50] longer shelf life. It is no longer on the market.

Improved nutrition
The GM oilseed crops on the market today offer improved oil profiles for processing or healthier edible [51] oils. The GM crops in development offer a wider array of environmental and consumer benefits such as nutritional enhancement and drought and stress tolerance. GM plants are being developed by both private companies and public research institutions such as CIMMYT, the International Maize and Wheat [52] Improvement Centre. Other examples include a genetically modified cassava with lower cyanogen [53] glucosides and enhanced with protein and other nutrients , while golden rice, developed by the International Rice Research Institute (IRRI), has been discussed as a possible cure for Vitamin A [54] deficiency. An international group of academics has generated a vitamin-enriched corn derived from South African white corn variety M37W with 169x increase in beta carotene, 6x the vitamin C and 2x [55] folate - it is not in production anywhere, but proves that this can be done.

Stress resistance
Plants engineered to tolerate non-biological stresses like drought, frost and nitrogen [61] [62] starvation or with increased nutritional value (e.g. Golden rice ) were in development in 2011.
[56][57] [58][59][60]

Herbicide resistance

Tobacco plants have been engineered to be resistant to the herbicide bromoxynil. And many crops have created that are resistant to the herbicide glyphosate. As weeds have grown resistant to glyphosate and other herbicides used in concert with resistant GM crops, companies are developing crops engineered to become resistant to multiple herbicides to allow farmers to use a mixed group of two, three, [64] or four different chemicals.

[63]

Pathogen resistance - insects or viruses

Tobacco, and may other crops, have been generated that express genes encoding for insecticidal [65] [66] proteins from Bacillus thuringiensis (Bt). Papaya, potatoes, and squash have been engineered to resist viral pathogens, such as cucumber mosaic virus which despite its name infects a wide variety of plants.

Production of biofuels
Algae, both hybrid and GM, is under development by several companies for the production of [67] biofuels. Jatropha has also been modified to improve its qualities for fuel product. SwissbasedSyngenta has received USDA approval to market a maize seed trademarked Enogen, which has been genetically modified to convert its own starch to sugar to speed the process of making ethanol for [68] biofuel.

Production of useful by-products

Drugs
Bananas have been developed, but are not in production, that produce human vaccines against infectious [69] diseases such as Hepatitis B. Tobacco plants have been developed and studied, but are not in [70] production, that can produce therapeutic antibodies.

Materials
Several companies and labs are working on engineering plants that can be used to make [71] [72] bioplastics. Potatoes that produce more industrially useful starches have been developed as well.

Bioremediation
Scientists at the University of York developed a weed (Arabidopsis thaliana) that contains genes from bacteria that can clean up TNT and RDX-explosive contaminants from the soil: It was hoped that this [73] weed would eliminate this pollution. 16 million hectares in the USA (1.5% of the total surface) are estimated to be contaminated with TNT and RDX. However the weed Arabidopsis thaliana was not tough enough to withstand the environment on military test grounds and research is continuing with the [74] University of Washington to develop a tougher native grass. Genetically modified plants have also been used for bioremediation of contaminated soils. Mercury, selenium and organic pollutants such as polychlorinated biphenyls (PCBs), TNT and RDXexplosive contaminants have been removed from soils by transgenic plants containing genes for bacterial enzymes.

Life Cycle of Gymnosperm and Angiosperm Gymnosperms Gymnosperms, like all vascular plants have a sporophyte-dominant life-cycle. The gametophyte (gametebearing phase) is relatively short-lived. Two spore types, microspores and megaspores, are typically produced in pollen cones or ovulate cones, respectively. Gametophytes, as with all heterosporous plants, develop within the spore wall. Pollen grains (microgametophytes) mature from microspores, and ultimately produce sperm cells. Megagametophytes develop from megaspores and are retained within the ovule. They typically produce multiple archegonia. During pollination, pollen grains are physically transferred between plants, from pollen cone to the ovule, being transferred by wind or insects. Whole grains enter each ovule through a microscopic gap in the ovule coat (integument) called the micropyle. The pollen grains mature further inside the ovule and produce sperm cells. Two main modes of fertilization are found in gymnosperms. Cycads andGinkgo have motile sperm that swim directly to the egg inside the ovule, whereas conifers and gnetophytes have sperm with no flagella that are conveyed to

the egg along a pollen tube. After fertilization (joining of the sperm and egg cell), the zygote develops into an embryo (young sporophyte). More than one embryo is usually initiated in each gymnosperm seed. The mature seed comprises the embryo and the remains of the female gametophyte, which serves as a food supply, and the seed coat (integument). Angiosperm Reproductive Flower Parts In general, angiosperms have a floral axis with four floral parts, two of which are fertile. At the receptacle, or tip, of the axis there is an ovule-bearing leaf structure known as the carpel. The ovule or ovules can be found inside the pistil. Three portions compose the pistil: the ovary, the style, and the stigma, where the pollen usually germinates. The mature ovule consists of the placenta, the integu- ments that are modified leaves that cover the entrance to the embryo sac, the micropyle, and the chalaza. These latter two parts of the ovule complement each other in their positions and functions. While the micropyle receives and guides the pollen tube, the chalaza relates to the vascular supply of the ovule, nutrition, and support. The stamens, which are often composed of the filament and sporangia sacs that make up the anther, surround the pistil. Stamens carry the male gametes, and the pistil carries the female gamete needed for sexual reproduction. It is believed that the great diversity and adaptability of the angiosperms is related to the presence of a unique reproductive cycle. This cycle consists of an alternation of generations and the production of a pair of spores on two types of sporophylls: microspores (which become male gametophytes) and megaspores (which become female gametophytes). Male Gamete Development The angiosperm reproductive cycle begins with the process of microsporogenesis, or microspore formation. The stamen consists of a filament and the anther, also known as the microsporangium. Inmost of the cases, the anther consists of four pollen sacs, or locules. Within each locule, the archesporial cell develops through mitosis and extends as a row of cells throughout the entire length of the young anther. These mitotic cell divisions generate the anther wall, which is made up of several cell layers, the outermost of which transforms itself into the epidermis. The layer of cells belowthe epidermis is known as the endothecium. During anther development, the endothecial cells acquire thickenings whose function is related to anther opening and pollen release. The innermost layer of the anther wall is the tapetum,whose primary function correlates with the nourishment of the young pollen and the deposition of the exine, a coating of the pollen grain. As development proceeds, the sporogenous cells located below the tapetum transform into microsporocytes. These microsporocytes will undergo meiosis, and tetrads (units of four) of microspores will form. Shortly after their formation, the tetrads separate into individual microspores. Each microspore is haploid, and often it will enlarge and separate from the tetrad, becoming sculptured by the deposition of sporopollenin and other substances that will turn into the ornamented surface of the pollen grain. The second phase of pollen development is known asmicrogametogenesis. Themicrospore is the first cell of the gametophytic generation, the cell that generates themature pollen. The single-nucleus microspore develops into the male gametophyte before the pollen is released.

This developmental process occurs through two or three unequalmitotic divisions of the nucleus and subsequent cytokinesis (cell separation). The two daughter nuclei and cells differ in size and in form. The larger cell represents the tube cell and nucleus,while the smaller cell represents the generative cell and nucleus. At maturity, the grain can be shred in two or three nucleate conditions. When the anther opens, the mature male gametophytes or pollen grains will be disseminated and ready for germination. Female Gamete Development The ovule (female sex organ) consists of two opposite ends: the micropyle, where the integuments come together, and a more distant end, where the ovular tissue is more massive. This part is also known as the chalaza, and it lies directly opposed to the micropyle. The mature ovule is composed of three layers: the outer integument; the inner integument; and, underneath the integuments, the nucellus. During ovular development, one cell lying below the nucellar epidermis changes into a primary archesporial; this will divide to form the primary parietal cell and primary sporogenous cell. The primary sporogenous cell functions as the megaspore mother cell, which divides meiotically, originating four haploid megaspores. In the majority of angiosperms, three of the megaspores will degenerate, and only the chalazal one will develop into the megagametophyte (embryo sac). After the completion of the embryo sac stage, a series of cellular events occurs, ending with the formation of the mature embryo sac, ready for fertilization by the male gametes. The chalazal megaspore enlarges and undergoes threemitoses, giving rise to eight haploid cells. The mature megagametophyte consists of two groups of four cells located at both ends of the embryo sac. The result is three antipodals at the chalazal end: the egg apparatus (consisting of the egg and two synergids at the micropylar end) and the polar nuclei. These two cells, present at both ends, usually fuse before pollination, and during fertilization they form the primary endosperm nucleus. Pollination The plant reproductive structures are now ready for the union of male and female gametes or fertilization, which eventually will produce a seed with a viable embryo and cotyledons. Before that step takes place, however, the pollen must be transferred from the anther to the stigma. Biotic agents (such as birds, insects, or mammals) or abiotic agents (such as wind or water) can accomplish this transfer process, known as pollination. After landing on the stigma, pollen tubes will emerge through the grain apertures if the environ- ment is high in humidity. Successful germination of the pollen in the stigma requires nutrients. In most plants, growth of the pollen tube lasts between twelve and forty-eight hours, frompollen germination to fertilization. Pollen germination starts with pollen-tube initiation, elongation, and penetration of the stigmatic tissue. During this period intense metabolic activity takes place, for the tube must synthesize membrane material and cell wall for growth and expansion. Simultaneously, at its tip the tube carries the vegetative cell nucleus, fol- lowed by the germinative cell. Angiosperms have evolved complex breeding systems that ensure theywill be pollinated by their own species. Today it is recognized that two pollination syndromes exist: self-pollination and cross-pollination. In self-breeding species, the pollen comes fromthe anther of the same flower. In cross-pollination (or outcrossing) species, the pollen comes from the anthers of a different flower or even a different plant of the same species. In these plants, incompatibility in the stigma guarantees that

only pollen from other flowers will germinate. Fertilization The union of one sperm with the egg is known as fertilization. However, several developmental processes in the vegetative and germinative cells prepare the two sperms for a process known as double fertilization. A mitotic division of the germinative cell generates the spermcells. This process that can take place on the growing pollen tube or inside the pollen grain. In a growing pollen tube, the vegetative nucleus disintegrates and the sperm cells will take the lead and enter the embryo sac for successful fertilization. Usually, the interactions between the pollen grain and the pistil ensure that the sperm cells will often reach the micropyle of the ovule. Once the spermreach themicropyle, the growth of other tubes stops. In the embryo sac (female gametophyte), four cells are located at themicropylar side.Of those four, the first pair that the spermcells will encounter are the synergids. One of these is always bigger than the other and carries the filiform apparatus, a structure resembling hairs that degenerates after pollination and before fertilization. The synergids act as chemical attractants to the pollen tube, which penetrates the synergids via the filiform apparatus and then releases the two sperm cells. One of the sperm cells will fuse with the egg, producing the zygote; the other sperm cell will fuse with the primary endosperm nucleus, generating the endosperm. The remaining cells of the female gametophyte are the antipodals; they usually degenerate after fertilization has taken place. Seed and Fruit Formation Once fertilization has occurred, the ovulewill go through a series ofmetabolic steps ending with the formation of the seed and the fruit. The recently created zygote transforms into amulticellular and complex embryo that has two well-defined polar ends: the radicle, or primary root, and the embryonic apical meristem with the first leaves. After successive mitosis, the mature endosperm usually grows close to the embryo and may provide nutrients needed for germination. The integuments will undergo further transformation, replication, and elongation and will become the seed coatof variable texture, consistency, and colors, depending on the type of plant. In general, after pollination or during fertilization, the ovary undergoes a series of physiological changes regulated by synchronized hormonal and genetic alterations that will modify the size of the parenchyma cells and its sugar and organic acids contents. This process turns the ovary into fruitin many cases familiar as the edible fruits familiar in human diets. The fruit provides nourishment for the seed until it ripens and drops to the ground, where the next stage in the life cycle begins. Germination, Seedling Development, and Maturation Seeds are released from the fruit in a large variety of ways that have evolved to ensure the survival of species. Whether ingested by mammals and passed through their feces to the ground, borne by wind on feathery "wings", or simply falling from rotting fruit that has abscissed and dropped from the plant, the seed must next undergo a process called germination, in which the embryo enclosed in the seed begins its growth. The embryo develops a hypocotyl (root axis) and a fleshy part known as the cotyledon; inmonocots there is one cotyledon, in dicots, two.

Germination requires certain conditions, such as the softening of the seed coat, moisture, and adequate warmth, to occur. During germination, the hypocotyl begins growing downward to become the root; the cotyledon(s) will develop into the shoot, stems, and leaves. The process of germination results in the sprouting through the grounds surface of the seedling, which will develop into the mature plant with flowers. The cycle then begins again.

Sporophyte All land plants, and some algae, have life cycles in which a haploid gametophyte generation alternates with a diploidsporophyte, the generation of a plant or algae that has a double set of chromosomes. A multicellular sporophyte generation or phase is present in the life cycle of all land plants and in some green algae. For common flowering plants (Angiosperms), the sporophyte generation makes up almost their whole life cycle (i.e. whole green plant, roots etc.), except phases of small reproductive structures [citation needed] (pollen and ovule). The sporophyte produces spores (hence the name), by meiosis. These meiospores develop into a gametophyte. Both the spores and the resulting gametophyte are haploid, meaning they only have one set of homologous chromosomes. The mature gametophyte produces male or female gametes (or both) by mitosis. The fusion of male and female gametes produces a diploidzygote which develops into a new sporophyte. This cycle is known as alternation of generations or alternation of phases.

In flowering plants, the sporophyte comprises the whole multicellular body except the pollen and embryo sac

In the normal course of events, the zygote and sporophyte will have a full double set of chromosomes again. An exception is when a diploid and haploid gamete fuse, resulting in a triploid sporophyte, which [citation will usually be sterile, as dividing three sets ofchromosomes into two halves causes complications.
needed]

Bryophytes (mosses, liverworts and hornworts) have a dominant gametophyte stage on which the adult sporophyte is dependent on the gametophyte for nutrition. The embryo of the sporophyte develops from the zygote within the female sex organ orarchegonium, and in its early development is therefore nurtured [1] by the gametophyte. Because this embryo-nurturing feature of the life cycle is common to all land plants they are known collectively as the Embryophytes.

Cleistocarpous sporophyte of the mossPhyscomitrella patens

Most algae have dominant gametophyte generations, but in some species the gametophytes and sporophytes are morphologically similar (isomorphic). An independent sporophyte is the dominant form in all clubmosses, horsetails, ferns,gymnosperms, and angiosperms (flowering plants) that have survived to the present day. Early land plants had sporophytes that produced identical spores (isosporous or homosporous) but the ancestors of the gymnosperms evolved complex heterosporous life cycles in which the spores producing male and female gametophytes were of different sizes, the female megaspores tending to be larger, and fewer in number, than the male microspores. During the Devonian period several plant groups independently evolved heterospory and subsequently the habit of endospory, in which single megaspores were retained within the sporangia of the parent sporophyte, instead of being freely liberated into the environment as in ancestral exosporous plants. These endosporic megaspores contained within them a miniature multicellular female gametophyte complete with female sex organs or archegonia containing oocytes which were fertilised by freeswimmingsperm produced by windborne miniatuarised male gametophytes in the form of pre-pollen. The resulting zygote developed into the next sporophyte generation while still retained within the pre-ovule, the single large female meiospore or megaspore contained in the modified sporangium or nucellus of the parent sporophyte. The evolution of heterospory and endospory were among the earliest steps in the evolution of seeds of the kind produced by gymnosperms and angiosperms today. Gametophyte A gametophyte is the haploid, multicellular phase of plants and algae that undergo alternation of generations, with each of its cells containing only a single set of chromosomes. The gametophyte produces male or female gametes (or both), by a process of cell division called mitosis. The female and male gametes are also called, respectively, egg cells and sperm. The fusion of male and female gametes produces a diploid zygote, which develops by repeated mitotic cell divisions into a multicellular sporophyte. Because the sporophyte is the product of the fusion of two haploid gametes, its cells are diploid, containing two sets of chromosomes. The mature sporophyte produces spores by a process called meiosis, sometimes referred to as "reduction division" because the chromosome pairs are separated once again to form single sets. The spores are therefore once again haploid and develop into a haploid gametophyte. In mosses, liverworts and hornworts (bryophytes), the gametophyte is the commonly known phase of the plant. An early developmental stage in the gametophyte of mosses (immediately following germination of the meiospore) is called the protonema. The adult gametophyte of mosses is called thegametophore as it [1] carries the gamete-producing sex organs. In most other land plants, the gametophyte is very small. In ferns the gametophyte is a free living [2] organism called the prothallus, in contrast to angiosperms. In gymnosperms and angiosperms, the gametophyte are reduced to only a few cells; in angiosperms the female gametophyte (sometimes called the "embryo sac") is known as a megagametophyte and the male gametophyte (pollen) is called a microgametophyte. In some multicellular green algae, red algae, or brown algae (Ulva is one example), the sporophytes and gametophytes are often isomorphic, but in some species the gametophyte may be reduced.

Plant Biological Clock A circadian clock, or circadian oscillator, is a biochemical mechanism that oscillates with a period of 24 hours and is coordinated with the day-night cycle. Circadian clocks are the central mechanisms which drive circadian rhythms. They consist of three major components: 1. A central oscillator with a period of about 24 hours that keeps time 2. A series of input pathways to this central oscillator to allow entrainment of the clock

3. A series of output pathways tied to distinct phases of the oscillator that regulate overt rhythms in biochemistry, physiology, and behavior throughout the organism The clock is reset as the environment changes through an organism's ability to sense external time cues of which the primary one is light. Circadian oscillators are ubiquitous in tissues of the body where they are synchronized by both endogenous and external signals to regulate transcriptional activity throughout the day in a tissue-specific manner.The basic molecular mechanisms of the biological clock have been defined in vertebrate species, Drosophila melanogaster, plants, fungi, and bacteria.

Plant hormones In general, it is accepted that there are five major classes of plant hormones, some of which are made up of many different chemicals that can vary in structure from one plant to the next. The chemicals are each grouped together into one of these classes based on their structural similarities and on their effects on plant physiology. Other plant hormones and growth regulators are not easily grouped into these classes; they exist naturally or are synthesized by humans or other organisms, including chemicals that inhibit plant growth or interrupt the physiological processes within plants. Each class has positive as well as inhibitory functions, and most often work in tandem with each other, with varying ratios of one or more interplaying to affect growth regulation. The five major classes are: [edit]Abscisic

acid

Abscisic acid also called ABA, was discovered and researched under two different names before its chemical properties were fully known, it was called dormin and abscicin II. Once it was determined that the two compounds are the same, it was named abscisic acid. The name "abscisic acid" was given because it was found in high concentrations in newly abscissed or freshly fallen leaves. This class of PGR is composed of one chemical compound normally produced in the leaves of plants, originating from chloroplasts, especially when plants are under stress. In general, it acts as an inhibitory chemical compound that affects bud growth, and seed and bud dormancy. It mediates changes within the apical meristem, causing bud dormancy and the alteration of the last set of leaves into protective bud covers. Since it was found in freshly abscissed leaves, it was thought to play a role in the processes of natural leaf drop, but further research has disproven this. In plant species from temperate parts of the world, it plays a role in leaf and seed dormancy by inhibiting growth, but, as it is dissipated from seeds or buds, growth begins. In other plants, as ABA levels decrease, growth then commences as gibberellin levels increase. Without ABA, buds and seeds would start to grow during warm periods in winter and be killed when it froze again. Since ABA dissipates slowly from the tissues and its effects take time to be offset by other plant hormones, there is a delay in physiological pathways that provide some protection from premature growth. It accumulates within seeds during fruit maturation, preventing seed germination within the fruit, or seed germination before winter. Abscisic acid's effects are degraded within plant tissues during cold temperatures or by its removal by water washing in out of the tissues, releasing [6] the seeds and buds from dormancy. In plants under water stress, ABA plays a role in closing the stomata. Soon after plants are waterstressed and the roots are deficient in water, a signal moves up to the leaves, causing the formation of ABA precursors there, which then move to the roots. The roots then release ABA, which is translocated to the foliage through the vascular system and modulates the potassium and sodium uptake within the guard cells, which then lose turgidity, closing the stomata. ABA exists in all parts of the plant and its concentration within any tissue seems to mediate its effects and function as a hormone; its degradation, or more properly catabolism, within the plant affects metabolic reactions and cellular growth and production of other hormones. Plants start life as a seed with high ABA levels. Just before the seed germinates, ABA levels decrease; during germination and early growth of the seedling, ABA levels decrease even more. As plants begin to produce shoots with fully functional leaves, ABA levels begin to increase, slowing down cellular growth in more "mature" areas of the plant. Stress from water or predation affects ABA production and catabolism rates, mediating another cascade of effects that trigger specific

responses from targeted cells. Scientists are still piecing together the complex interactions and effects of this and other phytohormones.

Auxins
The auxin indole-3-acetic acid

Auxins are compounds that positively influence cell enlargement, bud formation and root initiation. They also promote the production of other hormones and in conjunction with cytokinins, they control the growth [11] of stems, roots, and fruits, and convert stems into flowers. Auxins were the first class of growth [12] regulators discovered. They affect cell elongation by altering cell wall plasticity. They stimulate cambium, a subtype of meristem cells, to divide and in stems causesecondary xylem to differentiate. Auxins act to inhibit the growth of buds lower down the stems (apical dominance), and also to promote lateral and adventitious root development and growth. Leaf abscission is initiated by the growing point of a plant ceasing to produce auxins. Auxins in seeds regulate specific protein [13] synthesis, as they develop within the flower after pollination, causing the flower to develop a fruit to contain the developing seeds. Auxins are toxic to plants in large concentrations; they are most toxic to dicots and less so to monocots. Because of this property, synthetic auxin herbicides including 2,4D and 2,4,5-T have been developed and used for weed control. Auxins, especially 1-Naphthaleneacetic acid (NAA) and Indole-3-butyric acid (IBA), are also commonly applied to stimulate root growth when taking cuttings of plants. The most common auxin found in plants is indole-3-acetic acid or IAA. The correlation of auxins and cytokininsin the plants is a constant (A/C = const.).

Cytokinins
The cytokinin zeatin, Zea, in which it was first discovered in immature kernels.

Cytokinins or CKs are a group of chemicals that influence cell division and shoot formation. They were called kinins in the past when the first cytokinins were isolated from yeast cells. They also help delay senescence or the aging of tissues, are responsible for mediating auxin transport throughout the plant, and affect internodal length and leaf growth. They have a highly synergistic effect in concert with auxins, and the ratios of these two groups of plant hormones affect most major growth periods during a plant's lifetime. Cytokinins counter the apical dominance induced by auxins; they in conjunction with ethylene [14] promote abscission of leaves, flower parts, and fruits. The correlation of auxins and cytokinins in the plants is a constant (A/C = const.).

Ethylene
Ethylene is a gas that forms through the Yang Cycle from the breakdown of methionine, which is in all cells. Ethylene has very limited solubility in water and does not accumulate within the cell but diffuses out of the cell and escapes out of the plant. Its effectiveness as a plant hormone is dependent on its rate of production versus its rate of escaping into the atmosphere. Ethylene is produced at a faster rate in rapidly growing and dividing cells, especially in darkness. New growth and newly germinated seedlings produce more ethylene than can escape the plant, which leads to elevated amounts of ethylene, inhibiting leaf expansion (seeHyponastic response). As the new shoot is exposed to light, reactions by phytochrome in the plant's cells produce a signal for ethylene production to decrease, allowing leaf expansion. Ethylene affects cell growth and cell shape; when a growing shoot hits an obstacle while underground, ethylene production greatly increases, preventing cell elongation and causing the stem to swell. The resulting thicker stem can exert more pressure against the object impeding its path to the surface. If the shoot does not reach the surface and the ethylene stimulus becomes prolonged, it affects the stem's natural geotropic response, which is to grow upright, allowing it to grow around an object. Studies seem to indicate that ethylene affects stem diameter and height: When stems of trees are subjected to wind, causing lateral stress, greater ethylene production occurs, resulting in thicker, more sturdy tree trunks and branches. Ethylene affects fruit-ripening: Normally, when the seeds are mature, ethylene production increases and builds-up within the fruit, resulting in a climacteric event just before seed dispersal. The nuclear protein Ethylene Insensitive2 (EIN2) is regulated by ethylene production, and, in turn, regulates other hormones including ABA and stress hormones.

Gibberellins
Gibberellins, or GAs, include a large range of chemicals that are produced naturally within plants and by fungi. They were first discovered when Japanese researchers, including Eiichi Kurosawa, noticed a chemical produced by a fungus called Gibberella fujikuroi that produced abnormal growth in rice [16] plants. Gibberellins are important in seed germination, affecting enzyme production that mobilizes food production used for growth of new cells. This is done by modulating chromosomal transcription. In grain (rice, wheat, corn, etc.) seeds, a layer of cells called the aleurone layer wraps around the endosperm tissue. Absoption of water by the seed causes production of GA. The GA is transported to the aleurone layer, which responds by producing enzymes that break down stored food reserves within the endosperm, which are utilized by the growing seedling. GAs produce bolting of rosette-forming plants, increasing internodal length. They promote flowering, cellular division, and in seeds growth after germination. Gibberellins also reverse the inhibition of shoot growth and dormancy induced by ABA.

Seed Germination Germination is the growth of an embryonic plant contained within a seed; it results in the formation of the seedling. The seed of a vascular plant is a small package produced in a fruit or cone after the union of male and female sex cells. All fully developed seeds contain an embryo and, in most plant species some store of food reserves, wrapped in a seed coat. Some plants produce varying numbers of seeds that lack [1] embryos; these are called empty seeds and never germinate. Most seeds go through a period of dormancy where there is no active growth; during this time the seed can be safely transported to a new location and/or survive adverse climate conditions until circumstances are favorable for growth. Dormant seeds are ripe seeds that do not germinate because they are subject to external environmental conditions that prevent the initiation of metabolic processes and cell growth. Under favorable conditions, the seed begins to germinate and the embryonic tissues resume growth, developing towards a seedling. [edit]Factors

affecting seed germination

Seed germination depends on both internal and external conditions. The most important external factors [2] include temperature, water, oxygen and sometimes light or darkness. Various plants require different variables for successful seed germination. Often this depends on the individual seed variety and is closely linked to the ecological conditions of a plant's natural habitat. For some seeds, their future germination response is affected by environmental conditions during seed formation; most often these responses are types of seed dormancy. Water - is required for germination. Mature seeds are often extremely dry and need to take in significant amounts of water, relative to the dry weight of the seed, before cellular metabolism and growth can resume. Most seeds need enough water to moisten the seeds but not enough to soak them. The uptake of water by seeds is called imbibition, which leads to the swelling and the breaking of the seed coat. When seeds are formed, most plants store a food reserve with the seed, such as starch, proteins, or oils. This food reserve provides nourishment to the growing embryo. When the seed imbibes water, hydrolytic enzymes are activated which break down these stored food resources [2] into metabolically useful chemicals. After the seedling emerges from the seed coat and starts growing roots and leaves, the seedling's food reserves are typically exhausted; at this point photosynthesis provides the energy needed for continued growth and the seedling now requires a continuous supply of water, nutrients, and light. Oxygen - is required by the germinating seed for metabolism. Oxygen is used in aerobic [2] respiration, the main source of the seedling's energy until it grows leaves. Oxygen is an atmospheric gas that is found in soil pore spaces; if a seed is buried too deeply within the soil or the soil is waterlogged, the seed can be oxygen starved. Some seeds have impermeable seed coats that prevent oxygen from entering the seed, causing a type of physical dormancy which is broken when the seed coat is worn away enough to allow gas exchange and water uptake from the environment.
[3]

Temperature - affects cellular metabolic and growth rates. Seeds from different species and even seeds from the same plant germinate over a wide range of temperatures. Seeds often have a temperature range within which they will germinate, and they will not do so above or below this range. Many seeds germinate at temperatures slightly above 60-75 F (16-24 C) [room-temperature if you live in a centrally heated house], while others germinate just above freezing and others germinate only in response to alternations in temperature between warm and cool. Some seeds germinate when the soil is cool 28-40 F (-2 - 4 C), and some when the soil is warm 76-90 F (24-32 C). Some seeds require exposure to cold temperatures (vernalization) to break dormancy. Seeds in a dormant state will not germinate even if conditions are favorable. Seeds that are dependent on temperature to end dormancy have a type of physiological dormancy. For example, seeds requiring the cold of winter are inhibited from germinating until they take in water in the fall and experience cooler temperatures. Four degrees Celsius is cool enough to end dormancy for most cool dormant seeds, but some groups, especially within the family Ranunculaceae and others, need conditions cooler than -5 C. Some seeds will only germinate after hot temperatures during a forest fire which cracks their seed coats; this is a type of physical dormancy.

Most common annual vegetables have optimal germination temperatures between 75-90 F (24-32 C), though many species (e.g. radishes or spinach) can germinate at significantly lower temperatures, as low as 40 F (4 C), thus allowing them to be grown from seed in cooler climates. Suboptimal temperatures lead to lower success rates and longer germination periods. Light or darkness - can be an environmental trigger for germination and is a type of physiological dormancy. Most seeds are not affected by light or darkness, but many seeds, including species found in forest settings, will not germinate until an opening in the canopy allows sufficient light for growth of the seedling.

Scarification mimics natural processes that weaken the seed coat before germination. In nature, some seeds require particular conditions to germinate, such as the heat of a fire (e.g., many Australian native plants), or soaking in a body of water for a long period of time. Others need to be passed through an animal's digestive tract to weaken the seed coat enough to allow the seedling to emerge.

Dormancy
Some live seeds are dormant and need more time, and/or need to be subjected to specific environmental conditions before they will germinate.Seed dormancy can originate in different parts of the seed, for example, within the embryo; in other cases the seed coat is involved. Dormancy breaking often involves changes in membranes, initiated by dormancy-breaking signals. This generally occurs only within hydrated seeds. Factors affecting seed dormancy include the presence of certain plant hormones, notably abscisic acid, which inhibits germination, and gibberellin, which ends seed dormancy. In brewing, barley seeds are treated with gibberellin to ensure uniform seed germination for the production of barleymalt.

Seedling establishment
In some definitions, the appearance of the radicle marks the end of germination and the beginning of "establishment", a period that ends when the seedling has exhausted the food reserves stored in the seed. Germination and establishment as an independent organism are critical phases in the life of a plant [2] when they are the most vulnerable to injury, disease, and water stress. The germination index can be used as an indicator ofphytotoxicity in soils. The mortality between dispersal of seeds and completion of establishment can be so high that many species have adapted to produce huge numbers of seeds

Germination rate
In agriculture and gardening, the germination rate describes how many seeds of a particular plant species, variety or seedlot are likely to germinate. It is usually expressed as a percentage, e.g., an 85% germination rate indicates that about 85 out of 100 seeds will probably germinate under proper conditions. The germination rate is useful for calculating the seed requirements for a given area or desired number of plants.

Fruit Classification A fruit results from maturation of one or more flowers, and the gynoecium of the flower(s) forms all or part of the fruit. Inside the ovary/ovaries are one or more ovules where the megagametophyte contains the egg [10] cell. After double fertilization, these ovules will become seeds. The ovules are fertilized in a process that starts with pollination, which involves the movement of pollen from the stamens to the stigma of flowers. After pollination, a tube grows from the pollen through the stigma into the ovary to the ovule and two sperm are transferred from the pollen to the megagametophyte. Within the megagametophyte one of the two sperm unites with the egg, forming a zygote, and the second sperm enters the central cell forming the endosperm mother cell, which completes the double fertilization process. Later the zygote will give rise to the embryo of the seed, and the endosperm mother cell will give rise to endosperm, a nutritive tissue used by the embryo. As the ovules develop into seeds, the ovary begins to ripen and the ovary wall, the pericarp, may become fleshy (as in berries or drupes), or form a hard outer covering (as in nuts). In some multiseeded fruits, the extent to which the flesh develops is proportional to the number of fertilized ovules.The pericarp is often differentiated into two or three distinct layers called the exocarp (outer layer, also called epicarp), mesocarp (middle layer), and endocarp (inner layer). In some fruits, especially simple fruits derived from an inferior ovary, other parts of the flower (such as the floral tube, including the petals, sepals, and stamens), fuse with the ovary and ripen with it. In other cases, the sepals, petals and/or stamens and style of the flower fall off. When such other floral parts are a significant part of the fruit, it is called an accessory fruit. Since other parts of the flower may contribute to the structure of the fruit, it is important to study flower structure to understand how a particular fruit forms. There are three general modes of fruit development: Apocarpous fruits develop from a single flower having one or more separate carpels, and they are the simplest fruits. Syncarpous fruits develop from a single gynoecium having two or more carpels fused together. Multiple fruits form from many different flowers.

Plant scientists have grouped fruits into three main groups, simple fruits, aggregate fruits, and composite or multiple fruits.The groupings are not evolutionarily relevant, since many diverse plant taxa may be in the same group, but reflect how the flower organs are arranged and how the fruits develop.

Simple fruit

Epigynous berries are simple fleshy fruit. Clockwise from top right: cranberries,lingonberries, blueberries red huckleberries

Simple fruits can be either dry or fleshy, and result from the ripening of a simple or compound ovary in a flower with only one pistil. Dry fruits may be either dehiscent (opening to discharge seeds), or indehiscent [15] (not opening to discharge seeds). Types of dry, simple fruits, with examples of each, are: achene - Most commonly seen in aggregate fruits (e.g. strawberry) capsule (Brazil nut) caryopsis (wheat) Cypsela - An achene-like fruit derived from the individual florets in a capitulum (e.g. dandelion). fibrous drupe (coconut, walnut) follicle is formed from a single carpel, and opens by one suture (e.g. milkweed). More commonly seen in aggregate fruits (e.g. magnolia) legume (pea, bean, peanut) loment - a type of indehiscent legume nut (hazelnut, beech, oak acorn) samara (elm, ash, maple key)

schizocarp (carrot seed) silique (radish seed) silicle (shepherd's purse) utricle (beet)

Fruits in which part or all of the pericarp (fruit wall) is fleshy at maturity are simple fleshy fruits. Types of fleshy, simple fruits (with examples) are: berry (redcurrant, gooseberry, tomato, cranberry) stone fruit or drupe (plum, cherry, peach, apricot, olive)

An aggregate fruit, or etaerio, develops from a single flower with numerous simple pistils. Magnolia and Peony, collection of follicles developing from one flower. Sweet gum, collection of capsules. Sycamore, collection of achenes. Teasel, collection of cypsellas Tuliptree, collection of samaras.

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The pome fruits of the family Rosaceae, (including apples, pears, rosehips, and saskatoon berry) are a [17] syncarpous fleshy fruit, a simple fruit, developing from a half-inferior ovary. Schizocarp fruits form from a syncarpous ovary and do not really dehisce, but split into segments with [14] one or more seeds; they include a number of different forms from a wide range of families. Carrot seed is an example.

Aggregate fruit
Aggregate fruits form from single flowers that have multiple carpels which are not joined together, i.e. each pistil contains one carpel. Each pistil forms a fruitlet, and collectively the fruitlets are called an etaerio. Four types of aggregate fruits include etaerios of achenes, follicles, drupelets, and berries. Ranunculaceae species, including Clematis and Ranunculus have an etaerio of achenes, Calotropis has an etaerio of follicles, and Rubus species like raspberry, have an etaerio of drupelets.Annona have [18][19] Etaerio of berries. The raspberry, whose pistils are termed drupelets because each is like a small drupe attached to the receptacle. In some bramble fruits (such as blackberry) the receptacle is elongated and part of the ripe [20] fruit, making the blackberry an aggregate-accessory fruit. The strawberry is also an aggregate[21] accessory fruit, only one in which the seeds are contained in achenes. In all these examples, the fruit develops from a single flower with numerous pistils.

Multiple fruits
Main article: Multiple fruit A multiple fruit is one formed from a cluster of flowers (called an inflorescence). Each flower produces a [22] fruit, but these mature into a single mass. Examples are the pineapple, fig, mulberry, osage-orange, and breadfruit.
In some plants, such as this noni, flowers are produced regularly along the stem and it is possible to see together examples of flowering, fruit development, and fruit ripening.

In the photograph on the right, stages of flowering and fruit development in the noni or Indian mulberry (Morinda citrifolia) can be observed on a single branch. First an inflorescence of white flowers called a head is produced. After fertilization, each flower develops into a drupe, and as the drupes expand, they become connate (merge) into a multiple fleshy fruit called a syncarpet.

Berries
Berries are another type of fleshy fruit; they are simple fruit created from a single ovary. The ovary may be compound, with several carpels. Type include (examples follow in the table below): Pepo Berries where the skin is hardened, cucurbits Hesperidium Berries with a rind and a juicy interior, like most citrus fruit

Aggregate fruit
Aggregate fruit are formed from the merger of different ovaries of a single flower.

Accessory fruit
Some or all of the edible part of accessory fruit is not generated by the ovary.