a polyphyletic group and are placed in various divisions, some more closely related to plants than others. there are many differences between them in features such as cell wall composition, biochemistry, pigmentation, chloroplast structure and nutrient reserves. the algal division charophyta, sister to the green algal division chlorophyta, is considered to contain the ancestor of true plants. the charophyte class charophyceae and the land plant sub-kingdom embryophyta together form the monophyletic group or clade streptophytina. nonvascular land plants are embryophytes that lack the vascular tissues xylem and phloem. they include mosses, liverworts and hornworts. pteridophytic vascular plants with true xylem and phloem that reproduced by spores germinating into free-living gametophytes evolved during the silurian period and diversified into several lineages during the late silurian and early devonian. representatives of the lycopods have survived to the present day. by the end of the devonian period, several groups, including the lycopods, sphenophylls and progymnosperms, had independently evolved "megaspory" – their spores were of two distinct sizes, larger megaspores and smaller microspores. their reduced gametophytes developed from megaspores retained within the spore-producing organs (megasporangia) of the sporophyte, a condition known as endospory. seeds consist of an endosporic megasporangium surrounded by one or two sheathing layers (integuments). the young sporophyte develops within the seed, which on germination splits to release it. the earliest known seed plants date from the latest devonian famennian stage. following the evolution of the seed habit, seed plants diversified, giving rise to a number of now-extinct groups, including seed ferns, as well as the modern gymnosperms and angiosperms. gymnosperms produce "naked seeds" not fully enclosed in an ovary; modern representatives include conifers, cycads, ginkgo, and gnetales. angiosperms produce seeds enclosed in a structure such as a carpel or an ovary. ongoing research on the molecular phylogenetics of living plants appears to show that the angiosperms are a sister clade to the gymnosperms. plant physiology five of the key areas of study within plant physiology plant physiology encompasses all the internal chemical and physical activities of plants associated with life. chemicals obtained from the air, soil and water form the basis of all plant metabolism. the energy of sunlight, captured by oxygenic photosynthesis and released by cellular respiration, is the basis of almost all life. photoautotrophs, including all green plants, algae and cyanobacteria gather energy directly from sunlight by photosynthesis. heterotrophs including all animals, all fungi, all completely parasitic plants, and non-photosynthetic bacteria take in organic molecules produced by photoautotrophs and respire them or use them in the construction of cells and tissues. respiration is the oxidation of carbon compounds by breaking them down into simpler structures to release the energy they contain, essentially the opposite of photosynthesis. molecules are moved within plants by transport processes that operate at a variety of spatial scales. subcellular transport of ions, electrons and molecules such as water and enzymes occurs across cell membranes. minerals and water are transported from roots to other parts of the plant in the transpiration stream. diffusion, osmosis, and active transport and mass flow are all different ways transport can occur. examples of elements that plants need to transport are nitrogen, phosphorus, potassium, calcium, magnesium, and sulfur. in vascular plants, these elements are extracted from the soil as soluble ions by the roots and transported throughout the plant in the xylem. most of the elements required for plant nutrition come from the chemical breakdown of soil minerals. sucrose produced by photosynthesis is transported from the leaves to other parts of the plant in the phloem and plant hormones are transported by a variety of processes. plant hormones 1 an oat coleoptile with the sun overhead. auxin (pink) is evenly distributed in its tip. 2 with the sun at an angle and only shining on one side of the shoot, auxin moves to the opposite side and stimulates cell elongation there. 3 and 4 extra growth on that side causes the shoot to bend towards the sun. plants are not passive, but respond to external signals such as light, touch, and injury by moving or growing towards or away from the stimulus, as appropriate. tangible evidence of touch sensitivity is the almost instantaneous collapse of leaflets of mimosa pudica, the insect traps of venus flytrap and bladderworts, and the pollinia of orchids. the hypothesis that plant growth and development is coordinated by plant hormones or plant growth regulators first emerged in the late 19th century. darwin experimented on the movements of plant shoots and roots towards light and gravity, and concluded "it is hardly an exaggeration to say that the tip of the radicle . . acts like the brain of one of the lower animals . . directing the several movements". about the same time, the role of auxins (from the greek auxein, to grow) in control of plant growth was first outlined by the dutch scientist frits went. the first known auxin, indole-3-acetic acid (iaa), which promotes cell growth, was only isolated from plants about 50 years later. this compound mediates the tropic responses of shoots and roots towards light and gravity. the finding in 1939 that plant callus could be maintained in culture containing iaa, followed by the observation in 1947 that it could be induced to form roots and shoots by controlling the concentration of growth hormones were key steps in the development of plant biotechnology and genetic modification. cytokinins are a class of plant hormones named for their control of cell division (especially cytokinesis). the natural cytokinin zeatin was discovered in corn, zea mays, and is a derivative of the purine adenine. zeatin is produced in roots and transported to shoots in the xylem where it promotes cell division, bud development, and the greening of chloroplasts. the gibberelins, such as gibberelic acid are diterpenes synthesised from acetyl coa via the mevalonate pathway. they are involved in the promotion of germination and dormancy-breaking in seeds, in regulation of plant height by controlling stem elongation and the control of flowering. abscisic acid (aba) occurs in all land plants except liverworts, and is synthesised from carotenoids in the chloroplasts and other plastids. it inhibits cell division, promotes seed maturation, and dormancy, and promotes stomatal closure. it was so named because it was originally thought to control abscission. ethylene is a gaseous hormone that is produced in all higher plant tissues from methionine. it is now known to be the hormone that stimulates or regulates fruit ripening and abscission, and it, or the synthetic growth regulator ethephon which is rapidly metabolised to produce ethylene, are used on industrial scale to promote ripening of cotton, pineapples and other climacteric crops. another class of phytohormones is the jasmonates, first isolated from the oil of jasminum grandiflorum which regulates wound responses in plants by unblocking the expression of genes required in the systemic acquired resistance response to pathogen attack. in addition to being the primary energy source for plants, light functions as a signalling device, providing information to the plant, such as how much sunlight the plant receives each day. this can result in adaptive changes in a process known as photomorphogenesis. phytochromes are the photoreceptors in a plant that are sensitive to light. plant anatomy and morphology a nineteenth-century illustration showing the morphology of the roots, stems, leaves and flowers of the rice plant oryza sativa plant anatomy is the study of the structure of plant cells and tissues, whereas plant morphology is the study of their external form. all plants are multicellular eukaryotes, their dna stored in nuclei. the characteristic features of plant cells that distinguish them from those of animals and fungi include a primary cell wall composed of the polysaccharides cellulose, hemicellulose and pectin, larger vacuoles than in animal cells and the presence of plastids with unique photosynthetic and biosynthetic functions as in the chloroplasts. other plastids contain storage products such as starch (amyloplasts) or lipids (elaioplasts). uniquely, streptophyte cells and those of the green algal order trentepohliales divide by construction of a phragmoplast as a template for building a cell plate late in cell division. a diagram of a "typical" eudicot, the most common type of plant (three-fifths of all plant species). however, no plant actually looks exactly like this. the bodies of vascular plants including clubmosses, ferns and seed plants (gymnosperms and angiosperms) generally have aerial and subterranean subsystems. the shoots consist of stems bearing green photosynthesising leaves and reproductive structures. the underground vascularised roots bear root hairs at their tips and generally lack chlorophyll. non-vascular plants, the liverworts, hornworts and mosses do not produce ground-penetrating vascular roots and most of the plant participates in photosynthesis. the sporophyte generation is nonphotosynthetic in liverworts but may be able to contribute part of its energy needs by photosynthesis in mosses and hornworts. the root system and the shoot system are interdependent – the usually nonphotosynthetic root system depends on the shoot system for food, and the usually photosynthetic shoot system depends on water and minerals from the root system. cells in each system are capable of creating cells of the other and producing adventitious shoots or roots. stolons and tubers are examples of shoots that can grow roots. roots that spread out close to the surface, such as those of willows, can produce shoots and ultimately new plants. in the event that one of the systems is lost, the other can often regrow it. in fact it is possible to grow an entire plant from a single leaf, as is the case with plants in streptocarpus sect. saintpaulia, or even a single cell – which can dedifferentiate into a callus (a mass of unspecialised cells) that can grow into a new plant. in vascular plants, the xylem and phloem are the conductive tissues that transport resources between shoots and roots. roots are often adapted to store food such as sugars or starch, as in sugar beets and carrots. stems mainly provide support to the leaves and reproductive structures, but can store water in succulent plants such as cacti, food as in potato tubers, or reproduce vegetatively as in the stolons of strawberry plants or in the process of layering. leaves gather sunlight and carry out photosynthesis. large, flat, flexible, green leaves are called foliage leaves. gymnosperms, such as conifers, cycads, ginkgo, and gnetophytes are seed-producing plants with open seeds. angiosperms are seed-producing plants that produce flowers and have enclosed seeds. woody plants, such as azaleas and oaks, undergo a secondary growth phase resulting in two additional types of tissues: wood (secondary xylem) and bark (secondary phloem and cork). all gymnosperms and many angiosperms are woody plants. some plants reproduce sexually, some asexually, and some via both means. although reference to major morphological categories such as root, stem, leaf, and trichome are useful, one has to keep in mind that these categories are linked through intermediate forms so that a continuum between the categories results. furthermore, structures can be seen as processes, that is, process combinations. systematic botany a botanist preparing a plant specimen for mounting in the herbarium systematic botany is part of systematic biology, which is concerned with the range and diversity of organisms and their relationships, particularly as determined by their evolutionary history. it involves, or is related to, biological classification, scientific taxonomy and phylogenetics. biological classification is the method by which botanists group organisms into categories such as genera or species. biological classification is a form of scientific taxonomy. modern taxonomy is rooted in the work of carl linnaeus, who grouped species according to shared physical characteristics. these groupings have since been revised to align better with the darwinian principle of common descent – grouping organisms by ancestry rather than superficial characteristics. while scientists do not always agree on how to classify organisms, molecular phylogenetics, which uses dna sequences as data, has driven many recent revisions along evolutionary lines and is likely to continue to do so. the dominant classification system is called linnaean taxonomy. it includes ranks and binomial nomenclature. the nomenclature of botanical organisms is codified in the international code of nomenclature for algae, fungi, and plants (icn) and administered by the international botanical congress. kingdom plantae belongs to domain eukaryota and is broken down recursively until each species is separately classified. the order is: kingdom; phylum (or division); class; order; family; genus (plural genera); species. the scientific name of a plant