Angiosperms

Mark W Chase, Royal Botanic Gardens, Kew, UK

The angiosperms or flowering plants are the largest group of terrestrial, photosynthetic organisms, with perhaps 250 000 300 000 species occurring on all continents, including within the Arctic Circle and Antarctica.

Introductory article
Article Contents
. Introduction . Families and Classification . Economically Important Species . Horticulturally Important Species . Morphology

Introduction
The term angiosperm refers to the fact that the seeds of these plants are enclosed in several layers of tissue during their development, whereas those of the gymnosperms are exposed (naked). All angiosperms, regardless of whether they are herbaceous or woody, bear owers (although in some cases, these are so small that they go largely unnoticed). In many cases, the owers of these plants are conspicuous in their natural environments, and of course owers themselves are highly prized for their ornamental value. Flowering plants are the basis of life on Earth; they provide us with food, shelter, medicines and feed for our domesticated animals as well as ameliorating our environment by their production of oxygen and loss of water during photosynthesis (the cool shade under a tree on a hot day is a good example of the ability of plants to alter the local environment). In size, angiosperms range from less than 0.5 cm across (the smallest is Lepuropetalum from the Americas, whereas the largest are some of the tropical rainforest trees).

. Ecology . Biogeography . Fossil History . Phylogeny

name, Labiatae). Previous taxonomists have recognized two major groupings of owering plants based on the number of seed leaves (cotyledons) present in the seed and seedling stage, but the DNA sequence studies demonstrate that monocotyledons share unaperturate pollen with a group of dicotyledons (often termed the primitive dicotyledons), whereas the other dicots have triaperturate pollen. Thus the number of seed leaves is not a useful trait on which to split the angiosperms, whereas pollen features better reect the evolutionary history of these plants.

Economically Important Species

In economic terms, the most important food species are the cereals, which are all members of Poaceae (grasses); these include: rice (Oryza), wheat (Triticum), oats (Avena), corn or maize (Zea), and barley (Hordeum). Like all food species used by humankind, these species are all highly modied from the naturally occurring forms from which they were derived through domestication, and none of them would survive in nature outside of cultivation. Other important carbohydrate crops include yams (Dioscorea; Dioscoreaceae), yucca or cassava (Manihot; Euphorbiaceae), sweet potato (Ipomoea; Convolvulaceae), potato (Solanum; Solanaceae) and the several dierent groups of pulses (Cicer, Glycine, Lens, Phaseolus, Pisum and Vicia, which are chick pea, soya bean, lentil, beans, peas and broad bean, respectively; all of these are members of Fabaceae). Important vegetables include beet and chard (Beta; Amaranthaceae), cabbage, broccoli, cabbage, kale and cauliower (Brassica; Brassicaceae), carrot (Daucus; Apiaceae), aubergine and tomato (Solanum; Solanaceae), lettuce (Lactuca; Asteraceae), parsnip (Pastinaca; Apiaceae), peppers (Capsicum; Solanaceae), spinach (Spinacia; Amaranthaceae), squash, marrow and pumpkins (Cucurbita; Cucurbitaceae), and turnip and swede (Brassica; Brassicaceae). Among the fruits, the most important ones include apples (Malus; Rosaceae), avocado (Persea; Lauraceae),
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Families and Classification

The classication of the angiosperms has recently been the subject of some of the largest analyses of DNA sequence data ever conducted, both in terms of the number of genes included as well as the taxonomic scope of the sampling. There are roughly 450 families of owering plants according to these analyses, many of which have long histories of recognition, whereas some are only recently recognized, largely on the basis of the DNA studies. Only about ten families have not been studied in the DNA sequence analyses, and these are all small; most are rare as well. The largest of the angiosperm families and therefore most important in an evolutionary context are Asteraceae (the daisy or sunower family; an older name Compositae is often used by authors in Europe), Orchidaceae (orchid family), Fabaceae (bean family; an older name, Leguminosae is used frequently in Europe), Poaceae (grasses; Gramineae is an older name), Rubiaceae (coee family), Solanaceae (tomato and potato family), Euphorbiaceae (spurges and poinsettia), and Lamiaceae (mints; older

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cherries, plums and peaches (Prunus; Rosaceae), grapes (Vitis; Vitaceae), guava (Psidium; Myrtaceae), kiwifruit (Actinidia; Actinidiaceae), litchi (Litchi; Sapindaceae), mango (Mangifera; Anacardiaceae), melons (Cucumis; Cucurbitaceae), oranges and other citrus fruits (Citrus; Rutaceae), pears (Pyrus; Rosaceae), raspberries and blackberries (Rubus; Rosaceae), soursop and sweetsop (Annona; Annonaceae), and strawberries (Fragaria; Rosaceae). Spices and herbs include allspice (Pimenta; Myrtaceae), basil (Ocimum; Lamiaceae), cinnamon (Cinnamomum; Lauraceae), bay laurel (Laurus; Lauraceae), chocolate (Theobroma; Malvaceae), cloves (Syzygium; Myrtaceae), ginger (Zingiber; Zingiberaceae), nutmeg and mace (Myristica; Myristicaceae), oregano (Origanum; Lamiaceae), pepper (Piper; Piperaceae), rosemary (Rosmarinus; Lamiaceae) and vanilla (Vanilla; Orchidaceae). Sugar comes from at least two unrelated species, sugarbeet (Beta; Amaranthaceae) and sugarcane (Saccharum; Poaceae), as well as such things as maple (Acer; Sapindaceae). In addition to juices of many of the fruits listed above, (Ilex; beverages include coee (Coea; Rubiaceae), mate Aquifoliaceae) and tea (Camelia; Theaceae). Nuts come from a diversity of families, including almonds (Prunus; Rosaceae), chestnut (Castanea; Fagaceae), macadamia (Macadamia; Proteaceae), peanut (Arachis; Fabaceae), pecans (Carya; Juglandaceae), pistachio (Pistacia; Anacardiaceae) and walnut (Juglans; Juglandaceae). Housing materials and timber come from many families, such as ash (Fraxinus; Oleaceae), bamboo (Bambusa; Poaceae), beech (Fagus; Fagaceae), birch (Betula; Betulaceae), cherry (Prunus; Rosaceae), mahogany (Swietenia; Meliaceae), maple (Acer; Sapindaceae), oak (Quercus; Fagaceae), palms (many species of Arecaceae), teak (Tectona; Verbenaceae) and walnut (Juglans; Juglandaceae). Many other species of trees are used locally within dierent regions of the world, particularly in the tropics. Medicinal uses of plants are extensive and too numerous to mention here. Medical doctors in the Middle Ages were botanists and vice versa due to the fact that nearly all sources of medicines were plants, and to know which was which one had to be able to identify the plants of interest. Classication of plants was largely based on medical use, and such classications are still to be found among aboriginal peoples throughout the world. In modern times, the two professions, botanist and doctor, have been separated, but the use of plants as sources for medicinal compounds (which may later be synthesized in the laboratory) is an active and important area of research.

Our gardens are full of daodils (Narcissus; Amaryllidaceae), columbines (Aquilegia; Ranunculaceae), delphiniums (Delphinium; Ranunculaceae), gladioli (Gladiolus; Iridaceae), hibiscus (Hibiscus; Malvaceae), irises (Iris; Iridaceae), lilies (Lilium; Liliaceae), roses (Rosa; Rosaceae), sunowers (Helianthus; Asteraceae) and tulips (Tulipa; Liliaceae), but there are huge numbers of both wild species and hybrids in cultivation, and a complete list is not feasible here. Houseplants also include a great variety of species, such as African violets (Saintpaulia; Gesneriaceae), cacti (many genera of Cactaceae), orchids (many genera of Orchidaceae), palms (many genera of Arecaceae) and philodendron (Philodendron; Araceae).

Morphology
Flowering plants are the most diverse of all plants in the world; they can adapt to nearly all ecological conditions, and their vegetative features are as diverse as the areas they inhabit. They are composed of basically three organs: roots, shoots and leaves; having stated this, there are some plants (such as the bladderwort, Utricularia; Lentibulariaceae) that lack these distinctions and have just one type of organ that serves all purposes. Roots are largely of two types, thin, brous ones or thickened taproots; the rst type is purely absorptive in function whereas the second type can also serve a storage function (such as carrots and beets, Daucus and Beta). Many plants have a mixture of the two types, but some have primarily one or the other. Stems are likewise composed of two types: soft or herbaceous and woody. This dierence is due to the secondary (woody) growth that arises after the primary (nonwoody) stem has been produced. Secondary growth accounts for the increase in diameter of tree trunks over time, but some primary growth can be strengthened with bres at the time the primary growth is laid down, so that a great deal of woodiness can be produced without the need for secondary growth. Wood of trees is hard and resistant to decomposition, which is the basis of forestry and the logging industry. The various kinds of wood have dierent properties, and these dierences are reected in the underlying structure and organization of the cells making up the wood. The function of stems is support for and positioning of the photosynthetic organs, leaves, as well as the reproductive form of leaves, which are owers (see below). Stems are also modied for a number of storage functions, such as tubers (potatoes, Solanum, and dahlias, Dahlia, are a good example of this), bulbs (which are shorted stems covered by eshy leaf bases, as in tulips and lilies), and corms (as in crocus and gladiolus). These structures permit the plants to be dormant during dry or cold periods of the year when their existence above ground is not possible. Leaves are the most diverse of plant organs, and the dierences in shape and form represent modications to the

Horticulturally Important Species

In terms of horticulture, owering plants produce an amazing diversity of both garden plants and houseplants.
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ecological conditions under which the plant grows. The fundamental function of leaves is photosynthesis, but carrying out this function depends on the amount of water that is available in a given environment. A great deal of water is lost when plants are photosynthesizing because to facilitate gas exchange (the uptake of carbon dioxide and the production of oxygen), plant tissues must be in close contact with the atmosphere, which means that water in the plants tissues is lost to the environment. This is the reason why trees provide a cool place on a hot day; evaporation of water from their tissues absorbs heat and therefore cools the air. In habitats in which water is not freely available, plants must modify their leaves to counteract the eects of desiccation during photosynthesis, which is accomplished by alterations of both form and physiology. In dry environments, plants have narrow leaves covered by hairs or other surface ornamentation that locally diminish air movement, thus reducing water loss; plants also have evolved altered patterns of photosynthesis that permit them to operate at times of the day when the eects of wind and sun as not so great such as at night (this is referred to as crassulacean acid metabolism, CAM, or C4 photosynthesis). Flowers are made of up to four functionally specialized parts that are thought to have been derived evolutionarily from leaves. Sepals are the most leaf-like, and in many angiosperms they are green and protect the other oral parts until the owers are ready to open. Some species, such as lilies, orchids and irises, have sepals that take on the colour and shape of the petals, such that it appears that there are no sepals. Petals are nearly always colourful and function in the attraction of pollinating animals (bees, wasps, ies, butteries, and birds and even bats). They may also be modied to serve as a landing platform for the pollinator or even to exclude animals that are not suitable for pollination (the snap of a snapdragon, for example, permits only bees of the proper size and behaviour to gain access to the nectar within). Stamens (or anthers) bear two or so pollen sacs on the end of a stalk or lament; pollen is eaten by many insects during the process of transfer to another ower, so it is both a reward and an attractant (in most cases some of the pollen escapes from the pollinator and ends up being transferred to another ower, which then pollinates that second ower). Most plants carry out both male and female functions, but some species place the sexes into separate owers on the same plant or have separately sexed plants (as in most animals). Although having both male and female functions in the same plant could result in self-fertilization, most plants avoid this by separating the timing of the male and female function or preventing self-fertilization by either mechanical or physiological means (the latter is self-incompatibility, which means that self-fertilization cannot occur because sperm produced by the pollen is not allowed to reach the eggs located within the ovary of the owers on the same plant that produced the pollen). Ovules produce the egg cells, and these are located with the carpels of the ower, which nearly universally

occupy the centre of the ower. These structures are often terminated by a sticky structure(s) to which pollen sticks, and once germinated the pollen tubes grow down to where the developing ovules are located. Sperm is then released into the ovule to fertilize the egg cell. From this process, an embryo is produced (as in nearly all organisms), but in owering plants the unit of dispersal is a seed, which contains an early stage of embryo development plus some form of food storage (starches, sugar and oils, as well as amino acids and other nutrients) and a seed coat, which can be variously modied to assist in dispersal (e.g. feathery hairs that permit the seed to be carried by the wind or barbed hairs that attach the seed to animals). Once deposited in a suitable site, germination of the seed takes place, and the process is repeated unless interrupted by the new plant being eaten by an animal or attacked by fungi or bacteria. Some plants have developed strategies by which they parasitize another plant or a fungus, and these plants often forgo photosynthesis and are unnoticed until their ower stems appear above ground.

Ecology
As stated above, owering plants live on every continent and exist under every set of environmental condition except hot springs and polar ice. There are marine owering plants that compete with algae for space on the ocean oor as well as some of the largest terrestrial organisms that have ever occurred (the giant redwoods of California, which are gymnosperms, are the largest plants currently in existence, but some of the tropical forest trees are nearly as large as these). Flowering plants are remarkable in the diversity of their modications for existence in the diverse habitat types that occur on Earth. Cacti (in the family Cactaceae) and the cactoid spurges (Euphorbiaceae) inhabit all but the driest of deserts, whereas other owering plants are either submerged or oating aquatics that do not closely resemble their close relatives on dry land. Pollination ecology of owering plants is also diverse. Abiotic vectors (wind and water) move around pollen of some groups, and in this respect angiosperms are like many of the other, more ancient groups of plants. However, the owers of angiosperms have enabled them to make use of animals as dispersers of their pollen, a phenomenon that is known to occur only in the cycads (a gymnosperm) outside of the owering plants. A wide variety of insects eect pollination (bees and butteries are probably the most common), but other vectors include birds, bats and other small mammals, such as pygmy opossum and some rodents. Although many owers oer rewards to their pollinators in the form of nectar, pollen and in a few cases various forms of chemical compounds, owers of other species mimic rewardoering owers and deceive their pollinators often enough to eect pollination. Orchids are well known for their
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exceedingly diverse modications to attract and make use of a wide range of pollinators, including some that deceive naive, newly emerged male wasps and bees into thinking that their owers are females of these species and trick them into attempting to mate with the ower, during which they carry out pollination. Most pollination mechanisms are more straightforward than this, however. Seed dispersal involves another set of strategies, again including abiotic as well as biotic means. Many plants have winged seeds or parachute-like structures that permit them to be carried on the wind, sometimes for considerable distances. Others have such small seeds that they are easily transported by wind without the need for any extensions of the seed coat. Rivers and the oceans are exploited by other species, and the seed coat of such plants is largely impervious to water until specic events occur that initiate germination once the seed has landed in a suitable habitat. As mentioned above, many other types of seeds have barbs and other extensions that attach them to the skin or fur of animals, and these hitchhike a ride. Burrs and sticktights are two examples of these sorts of seeds, and in some cases it is the whole ovary that acts as the dispersal agent, with the seeds falling out slowly over time. In quite a number of cases, seeds are not freely released from the ovary of the mother plant, and a eshy fruit develops that is attractive to animals. These fruits along with their seeds are eaten by animals, and either the seeds are spat out (often the case with humans) or they pass through the digestive track of the animal without harm and are deposited in the faeces of the animal at some distance from the mother plant. Some seeds will not germinate until they receive such treatment. Recently it has been discovered that the seeds and dry fruits of some species require a re before either seed dispersal or germination can take place; in fact, some seeds will only germinate after they have been exposed to smoke (it is not the heat of the re that initiates readiness for germination, but rather the compounds that occur in smoke; in this case articial germination can be accomplished by watering the seeds with smoked water, i.e. water that has had smoke bubbled through it). Although re seems to most people to be an undesirable ecological event, many plants throughout the world are not only adapted to deal with res they require it to complete their life cycles. Although there is not sucient space here to include more information on plant ecology, it should be clear from these few examples that owering plants have developed a wide array of mechanisms for dealing with the physical world, and forces that we would otherwise consider destructive, such as res and other disturbances, play an important role in maintenance of this diversity.

known that certain families characterize particular habitats and continental areas; for example, cacti (Cactaceae) are prominent in the arid regions of the Americas but are absent from African and Asian deserts. Many areas in the southern hemisphere (which was not as drastically aected by episodes of glaciation in the past as the northern hemisphere) have highly diverse and localized taxa (species, genera and even families, such as the Protea family, Proteaceae). Some of these geographically restricted groups have evolved only recently, at a time when dispersal to other continents was not possible (this is probably the case with the cacti), but in other instances it is not clear why particular species or families are now restricted to only a small region of the globe. These taxa are known to be old enough to have become more widely distributed when the continental plates were much closer than they are today. There are a number of botanists who are actively involved with the study of plant geography because understanding these distributions is the key to a better understanding of past geological events. Tropical parts of the world have not experienced episodes of glaciation, but this does not mean that these regions have been characterized by environmental stasis. They have been subjected to periods of shifting aridity, and few regions of the tropics have had a consistently wet and stable history. However, it is also clear that at any given point in the last two hundred million years (the period during which owering plants and their ancestors are known to have occurred; see below), there have always been at least some regions on each of the continents that have supported vegetation adapted to warm and moist conditions. Much of the north temperate zone has been repeatedly stripped clean of vegetation by glaciers, whereas the temperate parts of the southern hemisphere have been little aected by this phenomenon (the southern hemisphere has a much greater portion of its surface covered by water, which has a moderating eect on cold temperatures, plus the ocean currents have carried warm water to the shores of even Antarctica until roughly 20 million years ago (Ma), so the current deep-freeze of the South Pole has not been characteristic of that continent throughout most of geological history). Thus, the southern hemisphere is rich in sets of relict species and families that although known to have been more broadly distributed in the past are now restricted to narrow distributions.

Fossil History
Angiosperms are well represented in the fossil record from about 120 Ma (the mid-Cretaceous period) until the present, but many of the earliest forms are diverse and recognizable as members of extant families, so it is clear that earlier forms had to have occurred. As eorts to reconstruct angiosperm phylogenetic relationships have

Biogeography
Flowering plants occur on all continents, and although there are not many in the harshest deserts and on Antarctica, some do manage to occur there. It is well
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Figure 1 Summary of phylogentic relationships for angiosperms inferred from analysis of rbcL, atpB and 18S rDNA sequences; the jackknife consensus tree (for groups receiving 4 50% support) is shown. The number of species in each clade is given in parentheses. Jackknife support is given below branches. From Soltis et al. (1999).

Gymnosperms

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Fagales (9) Curcurbitales (7) Rosales (18) Fabales (6) Zygophyllales Celastrales (8) Oxalidales (9) Malpighiales (44) Sapindales (13) Malvales (16) Brassicales (14) Crossosomatales (3) Myrtales (10) Geraniales (7) Saxifragales (25) Lamiales (34) Solanales (10) Gentianales (10) Garryales (4) Asterales (18) Dipsacales (7) Apiales (8) Aquifoliales (5) Cornales (13) Ericales (32) Myrothamnaceae/Gunneraceae (2) Berberidopsidales (2) Santalales (7) Caryophyllales (24) Buxaceae/Didymelaceae (3) Trochodendraceae (2) Proteales (4) Sabiaceae (2) Ranunculates (19) Ceratophyllales (1) Monocots Winterales Laurales (9) Magnoliales (10) Chloranthales (3) Piperales (8) Illiciaceae (1) Schisandraceae (1) Austrobaileyaceae (1) Nymphaeaceae (1) Amborellaceae (1) Taxaceae (1) Podocarpaceae (1) Pinaceae (1) Ginkgoaceae (1) Ephedraceae (1) Gnetaceae (1) Welwitschiaceae (1)

Rosids Magnoliids Asterids Eudicots

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progressed and increasingly revealed a consistent and robust pattern for basal angiosperms (Figure 1), palaeobotanists have refocused their eorts to look for fossils that accord with the taxa shown to be close to the base of the owering plants. This eort appears to have paid o, and several recent papers have focused on earlier than previously known members of various groups, again leading to the conclusion that the age of the angiosperms is older than that previously hypothesized. If a molecular clock approach is applied, a similar conclusion has been reached, although some of these molecular estimates are much older than expected (some as old as 350 Ma for the split between monocots and dicots, which seems an unlikely old age). Gymnosperms are the closest relatives of the angiosperms, and the extant members of this group are sister to the angiosperms (which means that both are monophyletic, i.e. each is more closely related to members of their own group than they are to any of the other group). Seed plants are known from the fossils that are at least 350 Ma old (dating from the Upper Carboniferous to the Lower Devonian). If the group of seed plants from which the angiosperms arose was one of the earliest to diverge from the seed plant lineage, then the date for the split between extant gymnosperms and angiosperms could be as old as 350 Ma. This is not to say that the lineage leading to angiosperms would have looked unlike other seed-bearing plants of that time; the traits that we associate with being an angiosperm could have arisen much later, but the lineage itself could thus be much older and not be distinguishable from other fossil seed plants. Current understanding of the fossil record may be limited by what scientists are searching for, and if they are not using the correct search image, then earlier members of the angiosperm line will not be discovered. The refocusing of research caused by angiosperm phylogenetic studies is a good case to illustrate how new discoveries has been aided by a clearer idea of what is expected to be present in the fossil record. Once the angiosperms appeared in the fossil record, they clearly quickly diversied such that by 90 Ma most of the modern families that we know today were in existence. This is congruent with the appearance in the fossil record of most of the modern groups of insects that act as pollinators of these plants today. Nearly all orders of angiosperms have at least one fossil representative from around 90 Ma, except for some of the putatively most recently evolved, such as Asterales, Lamiales and Poales (Figure 1).

Phylogeny
Over the past ten years the angiosperms have been the focus of a great deal of research on their phylogenetic relationships, particularly using DNA sequences from various genes in all three compartments of plant genomes (mitochondrial, plastid and nuclear). This has resulted in a clear picture of most of higher order relationships within the group. An unusual genus, Amborella, with a single species native to New Caledonia is alone sister to the rest of the owering plants, and this is followed in branching order by Nymphaeaceae and then a group composed of Austrobaileyaceae, Illiciaceae and Schisandraceae. A larger group composed of several orders, Laurales, Magnoliales, Piperales and Winterales, contains the majority of what have been referred to in the past as the ranalean complex, but their relationships to the eudicots, monocots and a couple of small families (Ceratophyllaceae and Chloranthaceae) are not yet clear. An ordinal classication of the families of owering plants was published in 1998, which was the rst time that a major group of organisms was reclassied based mostly on DNA evidence.

Further Reading
Chase MW and Fay MF (2001) Ancient owering plants: DNA sequences and angiosperm classication. Genome Biology 2: 1012.11012.4. Erhlich PR and Roughgarden J (1986) The Science of Ecology. New York: Macmillan. Foster AS and Giord EM (1974) Comparative Morphology of Vascular Plants. San Francisco: Freeman. Friis EM, Chaloner WG and Crane PR (eds) (1987) The Origins of Angiosperms and Their Biological Consequences. Cambridge: Cambridge University Press. Givnish TJ (ed.) (1986) On the Economy of Plant Form and Function. Cambridge: Cambridge University Press. Heiser CB (1981) Seed to Civilization: The Story of Food, 2nd edn. San Francisco: Freeman. Judd WS, Campbell CS, Kellogg EA and Stevens PF (1999) Plant Systematics: A Phylogenetic Approach. Sunderland, MA: Sinauer. Raven PH, Evert RF and Eichhorn SE (1999) Biology of Plants, 6th edn. San Francisco: Freeman. Richards AJ (1986) Plant Breeding Systems, 2nd edn. London: Chapman and Hall. Soltis PS, Soltis DE and Chase MW (1999) Angiosperm phylogeny inferred from multiple genes: a research tool for comparative biology. Nature 402: 402404.

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