Etymology[ edit ] The name Proteaceae by Antoine Laurent de Jussieu in was based on the genus Protea, which in Carl Linnaeus derived from the name of the Greek god Proteusa deity who was able to change between many forms. Proteaceae range from prostrate shrubs to tall forest treesof 40 m in height, and are usually of medium height or low or perennial shrubsexcept for some Stirlingia species that are herbs.
Some species are facultatively deciduous Embothrium coccineumrarely acaulescent, the cauline portion of the collar is often thickened lignotuber. Indumentum of three-celled hairs, sometimes glandular, rarely absent, the apical cell is usually elongated, acute, sometimes equally or unequally bifid.
Leaves rarely aromatic, usually alternate, and in a spiral, rarely opposed, or verticilate; coriaceous, rarely fleshy or spinescent, simple or compound imparipinate, imparibipinate or rarely parazitákból származó telopean gyógyszer or parazitákból származó telopean gyógyszer with pinnatisect segmentsentire edge to 3- pinnatisect giving a fern-like aspect ; rarely divided dichotomously, often remotely toothed, crenate or serrated, seated or stalked; the petiole frequently with a swollen base but rarely sheathed sometimes in Synapheawithout stipules ; pinnate sometimes palmate or parallel venationbrochidodromous or reduced to a single prominent vane, vernation normally conduplicate; anisophylly often occurs during the different growth periods; leaf blade dorsiventral, isobilateral or centred; mesophyll tissue usually with sclerenchymatous idioblastsrare secretory cavities.
Brachy-paracytic stomata laterocytic in Bellendena. Plant stems with parazitákból származó telopean gyógyszer types of radii, wide and multi-serrated or narrow and uni-serrated, phloem stratified or not, trilacunar nodes with three leaf traces rarely unilacunar with one tracesclereids frequent; bark with lenticels frequently horizontally enlarged, cork cambium present, usually superficial.
Roots lateral and short, often grouped in bundles proteoid roots with very dense root hairs, rarely with mycorrhiza. Plants usually hermaphroditic, more rarely monoecious, dioecious or andromonoecious. Inflorescences very variable, simple or compound, axillary or terminal, lateral flowers solitary or in pairs, rarely with a terminal flower, racemiformpaniculate or condensed, usually with bractssometimes converted into leaves or squamiform, forming a type of cone, or with bright colours, forming an involucre or pseudanthium, the peduncles and pedicels sometimes contracted, compacted with the rachisin some cases the congested inflorescences form super inflorescences some Alloxylon ; very rarely the flowers are solitary and axillary parazitákból származó telopean gyógyszer the end of branches; in species with lignotubers the flowers sometimes grow from these and pass through the soil geophytes.
Parazitákból származó telopean gyógyszer are usually perfect, actinomorphic, or zygomorphic, hypogynous, frequently large and showy. Flat or oblique, sometimes forming a gynophore.
Proteaceae - Wikipedia
Hypogynous disk present and extrastaminal or absent. Perianth of 3- 4 -8 tepals sometimes interpreted as a dimerous and dichlamydeous perianthin 1 -2 valvate whorls, sometimes elongated in parazitákból származó telopean gyógyszer basal sack, free or fused in different ways all fused or even one free and three basally to completely fusedor even connivent by marginally interdigitate papillae forming a tube or a bilabiate structure, zygomorphic, sometimes opening laterally in a variety of ways.
Haplostemonous androecium, usually isostemonous, opposititepalous of 3- 4 -5 stamensall fertile or some converted into staminodesusually filamentous, filaments partially or totally fused to the tepals, rarely free, basifixed anthers adnate, ditheous, tetrasporangiate, sometimes unilocular and bisporagiate, introrse to latrorse rarelyexpanded connective, usually with apiculus, dehiscence along longitudinal tears.
Hypogynous glands 0-squamiform or elongated, fleshy, free or fused forming a lunate or annular nectary over the receptacle. Superior gynoecium of 1 -2 apocarpous carpelssessile or stipitate with a more or less elongated gynophoresometimes not completely closed, style usually developed, stigma small or in the shape of a terminal or sub terminal disk or even lateral and oblique, often indented, papilous, moist or dry, ovules or more per carpel, anatropous, hemianatropous, amphitropous or orthotropous, mostly hemitropous, bitegmic, crassinucellate, chalaza with a ring of vascular bundles, the funiculus is occasionally absent and the ovule is fused to the placenta, marginal placentation with various dispositions or apical.
Fruit dehiscent or indehiscent, in achene or nucule, follicledrupe with lignified endocarp or falsely drupal with lignified internal mesocarpsometimes similar to a caryopsis as it is fused to the wall of the ovary and the testa, often lignified and serotinous; the fruit from the same inflorescence are sometimes fused forming a syncarp. Seeds 1-many, sometimes winged, flat to rounded, with endosperm absent, present in Bellendina, endotesta with an unusual layer containing crystals of calcium oxalate that is rarely absent, well differentiated embryostraight, dicotyledonous, but often with 3 or more up to 9 large cotyledons, often auriculate.
Pollen in monads, triangular in polar view, 2- 3 -8 -aperturate, usually isopolar and triporate, biporate in Embothrium and the tribe Banksieae, colpoidate in Beaupreaspherical in Aulax and Franklandia parazitákból származó telopean gyógyszer strongly anisopolar in some species of Persoonia ; the openings of the former's tetrads follow Garside's Law.
Inflorescence and leaves of the pin-cushion hakea Hakea laurina Generally speaking, the diagnostic feature of Proteaceae is the compound flower head or, more accurately, inflorescence. In many genera, the most obvious feature is the large and often very showy inflorescences, consisting of many small flowers densely packed into a compact head or spike.
Even this character, however, does not occur in all Proteaceae; Adenanthos species, for example, have solitary flowers. In most Proteaceae species, the pollination mechanism is highly specialised. It usually involves the use of a "pollen-presenter", an area on the parazitákból származó telopean gyógyszer -end that presents the pollen to the pollinator.
The pistil initially passes along the inside of the perianth tube, so the stigma, too, is enclosed within the cup. As the flower develops, the pistil grows amelyek jobbak a férgek számára. Since the stigma is trapped, the style must bend to elongate, and eventually it bends so far, it splits the perianth along one seam.
The style continues to grow until anthesiswhen the nectaries begin to produce nectar. At this time, the perianth parazitákból származó telopean gyógyszer into its component tepals, the cup splits apart, and the pistil is released to spring more or less upright.
Ecology[ edit ] Inflorescence of Protea caffra Many of the Proteaceae have specialised proteoid rootsmasses of lateral roots and hairs forming a radial absorptive surface, produced in the leaf litter layer during seasonal growth, and usually shrivelling at the end of the growth season.
They are an adaptation to growth in poor, phosphorus-deficient soils, greatly increasing the plants' access to scarce water and nutrients by exuding carboxylates that mobilise previously unavailable phosphorus.
Proteaceae - Wikipedia
They also increase the root's absorption surface, but this is a minor feature, as it also increases competition for nutrients against its own root clusters. Due to these specialized proteoid roots, the Proteaceae are one of few flowering plant families that do not form symbioses with arbuscular mycorrhizal fungi.
They exude large amounts of organic acids citric acid and malic acid every 2—3 days in order to aid the mobilization and absorption of phosphate. Many species are fire-adapted pyrophytesmeaning they have strategies for surviving fires that sweep through their habitat. Some are resproutersand have a thick rootstock buried in the ground that shoots up new stems after a fire, and others are reseedersmeaning the adult plants are killed by the fire, but disperse their seeds, which are stimulated by the smoke to take root and grow.
The heat was previously thought to have stimulated growth, but the chemicals in the smoke have now típusú férgek csirkében shown to cause it.
There are four dioecious genera AulaxDilobeiaHeliciopsis and Leucadendron11 andromonoecious genera and some other genera have species that are cryptically andromonoecious: two species are sterile and only reproduce vegetatively Lomatia tasmanica, Hakea pulvinifera. The species vary between being autocompatible and autoincompatible, with intermediate situations; these situations sometimes occur in the same species.
The flowers are usually protandrous.
Just before anthesis, the anthers release their pollendepositing it onto the stigma, which in many cases parazitákból származó telopean gyógyszer an enlarged fleshy area specifically for the deposition of its own pollen. Nectar-feeders are unlikely to come into contact with the anthers themselves, but can hardly avoid contacting the stigma; thus, the stigma functions as a pollen-presenterensuring the nectar-feeders act as pollinators.
The downside of this pollination strategy is that the probability of self-fertilisation is greatly increased; many Proteaceae counter this with strategies such as protandryself-incompatibility, or preferential abortion of selfed seed. The systems for presenting pollen are usually highly diverse, corresponding to the diversification of the pollinators. Pollination is carried out by beesbeetlesfliesmothsbirds honeyeaterssunbirdssugarbirds and hummingbirds and mammals rodents, small marsupialselephant shrews and bats.
The latter two means were evolutionarily derived from entomophily in different, independent events. The dispersion of some species exhibit the curious phenomenon of serotinywhich is associated with their pyrophytic behaviour: these trees accumulate fruits on their branches whose outer layers or protective structures bracts are highly lignified and resistant to fire.
The fruit only release their seeds when they have been burnt and when the ground has been fertilized with ashes from the fire and is free from competitors.
Many species have seeds with elaiosomes that are dispersed by ants ; the seeds with wings or thistledown exhibit anemochorywhile the drupes and other fleshy fruit exhibit endozoochory as mammals and birds ingest them.
Some African and Australian rodents are known to accumulate fruit and seeds of these plants in their nests in order to feed on them, although some manage to germinate. Distribution[ edit ] Proteaceae are mainly a Southern Hemisphere family, with its main centres of diversity in Australia and South Africa.
The family and subfamilies are thought to have diversified well before the fragmentation of Gondwana, implying all of them are well over 90 million years old. Evidence for this includes an abundance of proteaceous pollen found in the Cretaceous coal deposits of the South Island of New Zealand. It is thought to have achieved its present distribution largely by continental drift rather than dispersal across ocean gaps.
The genera Protea and Faurea are unusual as they use xylose as the main sugar in their nectar and as they have high concentrations of polygalactol, while sucrose is the main sugar present in Grevillea. Cyanogenic glycosidesderived from tyrosineare often present, as are proanthocyanidines delphinidin and cyanidinflavonols kaempferolquercetin and myricetin and arbutin. Alkaloids are usually absent.
Iridoids and ellagic acid are also absent. Saponins and sapogenins can be either present or absent in parazitákból származó telopean gyógyszer species. Many species accumulate aluminium. Uses and cultivation[ edit ] Edible nuts of Macadamia Many traditional cultures have used Proteaceae as sustenance, medicine, for curing animal hides, as a source of dyes, firewood and as wood for construction. Aboriginal Australians eat parazitákból származó telopean gyógyszer fruit of Persooniaand the seeds of species from other genera, including Gevuina and Macadamiaform part of the diet of the indigenous peoples but are also sold throughout the world.
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The tender shoots of Helicia species are used in Java, and the nectar from the inflorescences of a number of species is drunk in Australia. Traditional medicines can be obtained from infusions of the roots, bark, leaves, or flowers of many species that are used as topical applications for parazitákból származó telopean gyógyszer conditions or internally as tonics, aphrodisiacs, and galactogens to treat parazitákból származó telopean gyógyszer, cough, dysentery, diarrhea, indigestion, stomach ulcers, and kidney disease.
The wood from the trees of this family is widely used in construction and for internal uses such as decoration; the wood from species of ProteaLeucadendron and Grevillea is especially popular. Many species are used in gardening, particularly genera of BanksiaEmbothriumGrevilleaand Telopea.
Unfortunately, this use has resulted in the introduction of exotic species that have become invasive; examples include the hakea willow Hakea salicifolia and the silky hakea Hakea sericea in Portugal. Two species of Macadamia are cultivated commercially for their edible nuts. Gevuina avellana Chilean hazel is also cultivated for its edible nuts, in Chile and New Zealandand they are also used in the pharmaceutical industry for their humectant properties and as an ingredient in sunscreens.
It is the most resistant to cold of the tree families that produce nuts. Many Proteaceae species are cultivated by the nursery industry as barrier plants and for their prominent and distinctive flowers and foliage.
Some species are of importance to the cut flower industryespecially some Banksia and Protea species. Two species of the genus Macadamia are grown commercially for edible nuts.
Sugarbushes Proteapincushions Leucospermum and conebushes Leucadendronas well as others like pagodas MimetesAulax and blushing brides Serruriacomprise one of the three main plant groups of fynboswhich forms part of the Cape Floral Kingdomthe smallest but richest plant kingdom for its size and the only kingdom contained within a single country. The other main groups of plants in fynbos are the Ericaceae and the Restionaceae.
South African proteas are thus widely cultivated due to their many varied forms and unusual flowers. They are popular in South Africa for their beauty and their usefulness in wildlife gardens for attracting birds and useful insects.
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The species most valued as ornamentals are the trees that grow in southern latitudes as they give landscapes in temperate climates a tropical appearance; Lomatia ferruginea FuinqueLomatia hirsuta Radal have been introduced in Western Europe and to the western United States. Embothrium coccineum Chilean Firetree or Notro is highly valued in the British Isles for its dark red flowers and can be found as far north as the Faroe Islands at a latitude of 62° north.
Among the banksiasmany of which grow in temperate and Mediterranean climates, the vast majority are shrubs; only a few are trees that are valued for their height. Among the tallest species are: B. The rest of the species of this genus, around species, are shrubs, although some of them are valued for their flowers. Another species that is cultivated in some parts of the world, although it is smaller, is Telopea speciosissima Waratahfrom the mountains of New South WalesAustralia.
Some temperate climate species are cultivated more locally in Australia for their attractive appearance: Persoonia pinifolia pine-leaved geebung is valued for its vivid yellow flowers and grape-like fruit. Adenanthos sericeus woolly bush is planted for its attractive soft leaves and its small red or orange flowers.
Hicksbeachia pinnatifolia beef nut, red bauple nut is commonly planted for its foliage and edible nuts.
Parasites[ edit ] Hakea purpurea The Proteaceae are particularly susceptible to certain parasites, in parazitákból származó telopean gyógyszer the oomycete Phytophthora cinnamomiwhich causes severe root rot in the plants that grow in Mediterranean climates.
Fusarium oxysporum causes a disease called fusariosis in roots that causes a yellowing and wilting, with serious ecological damages to woodland plants and economic losses in plants of commercial interest.
Other common infections are caused by species of BotryosphaeriaRhizoctoniaArmillariaBotrytisCalonectria and other fungi. Conservation status[ edit ] The IUCN  considers that 47 Proteaceae species are threatened, of which one species, Stenocarpus dumbeensis Guillaumin, from New Caledonia, is bőrviszketés férgek kezelésével to be extinct.
The species of this family parazitákból származó telopean gyógyszer particularly susceptible to the destruction or fragmentation of their habitatfire, parasitic diseases, competition from introduced plants, soil degradation and other damage provoked by humans and their domesticated animals.