Carnations also have unwanted pollinators. Those that do not fly, but crawl. Crawlers are disadvantageous in that they do not get from one plant to another and do not provide cross-pollination. To prevent crawling to the flowers, sticky protection is arranged just below the inflorescence.
The most reliable is the sticky defense of the drooping tar. Smolevka is a grass with single stems. The leaves, like those of other cloves, are attached oppositely. Two against each other, narrow, elongated. The flowers are long, like tubules, and only at the end unfold in a white fan. In loose inflorescence several flowers. Ants storm the sticky ring. After all, the inflorescence is at hand. Here it is, next to it. But it is impossible to overcome the adhesive barrier. Some stick and stay here, others retreat without success. Other cloves have the same sticky barriers: tar and white doze. But there are no barriers for flying pollinators. Smolevka is pollinated by moths: Therefore, the flowers are white. And the smell they have at night is strong, pleasant. During the day, the flowers close and become inconspicuous, because their underside is greenish.
Cloves seeds ripen in dry boxes. Neither birds nor animals are attracted. They don't have wings or parachutes. The calculation is different. The boxes are stretched upwards and have a hole at the very top. The hole is smaller than the diameter of the box itself. Seeds from the cloves are not blown out, but thrown out. Stems are high. When dry, they spring well. As soon as the wind blows, they begin to swing. But while the breeze is weak, the seeds only roll inside. Finally, a strong jerk of the wind, the stalk deviates to one side, then sharply back, and like a bullet, a seed flies out.
Peacock carnation. Photo: Ettore Balocchi
Even better, if the wind does not shake, but touches the animal. One of the gerbils, Sitnikova, has established a close relationship with domestic horses. This is a squat grass. Grows in turfs. The leaves are more like bristles than leaves. Sticking up from the spine in thick bunches. On bare stalks, a dozen large flowers. But horses are not interested in tops, but in roots. Especially when there is snow in the steppes where the gerbil grows.
In the steppe, the layer of snow is thin, bare turf is visible in places. Horses find gerbils quickly. They pound the frozen ground with their hooves, extracting roots. While hammering, the stalks with boxes are shaken more than once so that the seeds fly away in three meters. Then everything will turn into dust. She will scatter over the snow spots. It will fly even farther.
In the coastal part of the Canadian Arctic archipelago, botanist D. Saville noticed an even more curious phenomenon. In May, when the snow was still strong and smooth, like asphalt, the animals touched dry blades of grass, boxes broke off from them, and the wind, not even very strong, picked them up and drove them far, several hundred meters away. He believes that such a form of transport can carry boxes and seeds so far away that they move from one island of the archipelago to another.
Probably, this method of distribution also happens in the mountains. In the highlands, live clove plants can also move. True, it is slower and not everywhere, but in those places where stone taluses crawl along the slopes. The highlands are also the earth's outskirts. Above 6 thousand meters, flowering plants no longer grow. The height record here, as well as at the pole, is behind the cloves. It is they who are very often the first to settle on such unstable soil as scree. They slide down with her.
In 2012, Russian scientists published unusual news in the Bulletin of the US National Academy of Sciences - they returned from oblivion a small plant that lived in ice age.
The species, scientists were able to bring back to life after tens of thousands of years of being in permafrost, - narrow-leaved tar. From the clove family - widespread in Yakutia and now, being a local endemic. As now, and in the past, the narrow-leaved resin grew in the inhospitable conditions of the tundra and arctic deserts, on stony and sandy soils, often being one of the few species of flowering plants in these inhospitable conditions. By the way, representatives of the clove family are recognized cryophiles - the most resistant to frost - for example, one of the two species of Antarctic angiosperms - Kolobant Quito, belongs to this family.
Therefore, the prevalence of narrow-leaved resin in the ecosystems of the Ice Age was reflected in the diet of many animals, and in particular the American ground squirrel, in the fossil nests of the colonies of which scientists found significant frozen stocks of this grass. In frozen conglomerates of sand, plants and ice, which, according to radiocarbon analysis, are about 32 thousand years old, scientists found thousands of seeds - in some gopher nests there were 600-800 thousand of them. Most of these seeds, and the size of a separate one is several millimeters, were damaged by ground squirrels, however, this was enough and quite normal for the revival of the species. It was from these "normal" seeds that the work with the "resurrection" of the Ice Age plant took place.
In the Tien Shan and in the Sayan mountains, large, like anemones, flowers of the loosestrife sapling dazzlingly sparkle on the screes. The plant itself between the stones is barely noticeable, if not for the flowers. The stems are thin, shaggy from insulating hairs. One stalk between the stones cannot withstand. Therefore, the sapling grows in a heap, with a broom. Stones will press one stalk, cut it off. The bundle will survive. Creeping rhizomes creep under the stones. Directed more and more up the slope. The roots do not particularly tend to grow up. But the scree, sliding down, drags the bushes along with it. The roots of these bushes are firmly anchored in a layer of soil along which scree creeps. Under the load of piled stones, the roots are pulled down, the living parts are covered with rubble. New shoots grow from the roots, and new bushes grow on them down the slope. So step by step the bushes appear lower and lower down the slope. And the old roots are higher and higher in relation to them. This creates the illusion of roots growing upwards.
In the end, when there are a lot of bushes on the scree, they will begin to slow down the stone stream. And then other herbs settle on the slightly calmed scree. Among them, there are also cloves. More often gerbils. They slow down the movement of rubble even more. And finally, under their protection, the so-called dam plants settle, which finally pin the scree to the ground on which it travels.
Living on stones, the plant is forced to feed on what the stone gives. And involuntarily store up, accumulate in oneself what he is rich in. Some cloves accumulate so much tin, copper, selenium that they began to search for ore deposits. Geologists who conducted exploration in Rudny Altai noticed that on the sites of the old Demidov developments of copper ore, bushes of Patren's kachim are constantly found. Kachim is visible from afar: half a meter high, like a bouquet of shining pink flowers stuck into the ground. The stem is spirally twisted and goes along the cracks far into the depths of the rock. The leaves are gray, narrow, like all carnations, sit opposite.
We tried to find out how Patren's kachim is tied to copper ore. They put the outlines of ore deposits on the map. Then we marked the kachima bushes with dots. All points coincided with ore bodies. Now kachim has become a reliable indicator of copper ores.
Clove family (Caryophyllaceae) (V.V. Bochantseva, E.V. Semacheva)
Clove - one of the largest families in the order of clove. It contains approximately 80 genera and 2000 species. Carnations can be found on all continents of the globe, in a wide variety of habitats. Representatives of the family grow in the tundra, among the carnations there are many forest and meadow plants. They are also found in arid regions: in the steppes, semi-deserts and deserts. In the mountains, carnations rise to the alpine belt, and one of the species, creeping chickweed (Stellaria decumbens), was found in rocky crevices in the Himalayas at an altitude of 6000 m, much higher than other flowering plants of the highlands.
Carnations are especially widely represented in the temperate regions of the northern hemisphere, with the largest number of genera and species concentrated in the Mediterranean, Western and Central Asia. In the composition of the floras of most parts of the Arctic, representatives of this family
most often occupy the fifth place in terms of the number of species. Several genera of cloves are found in the southern hemisphere, in its temperate zone, as well as in the mountains of the tropical region. Even on ice-covered Antarctica, where there are practically no higher plants, on the Antarctic Peninsula of the mainland, along with the Antarctic pike grass, one species of clove was found - Colobanthus quitensis. A significant number of species and even genera of cloves are cosmopolitans. Such are the genera toritsa (Spergula), morinka (Spergularia), chickweed (Stellaria), chickweed (Cerastium), gerbil (Arenaria), etc.
The whole variety of genera and species of cloves, in accordance with the family system proposed by F. Paxo.m and K. Hoffman, is usually divided into three subfamilies: paronychia (Paronychioideae), alsin (Alsinoideae) and smolevkovye (Silenoideae); representatives of the subfamilies differ in the presence or absence of stipules and some structural features of the flowers.
The leaves of the carnations are opposite, rarely alternate, simple, entire, often narrow - linear or linear-lanceolate, equipped with scaly stipules (most paronychia) or without stipules (alsinovye and resinous). The flowers are usually collected in dichasial inflorescences very characteristic of this family, either highly branched and loose (in Kachima species - Gypsophila), or more compact, umbellate or corymbose (bearded carnation - Dianthus barbatus); much less often single flowers (common cockle - Agrostemma githago). The flowers are actinomorphic, in most representatives 5-membered (Fig. 199). Sepals 5, free, or almost free (paronychia and alsinovye) or fused into a tube (small, for example, firecracker - Silene vulgaris, pl. 55); often there are bracts close to the calyx. Petals usually 5, always free; in some species, the petals are barely developed or completely absent. Only in resinous petals have long nails (the narrowed part of the petals) and limbs whole or divided into narrow lobes (the expanded part of the petals), and on the border between them sometimes there are petal-like outgrowths forming the so-called accessory corolla or corolla. Stamens 10 arranged in two circles or 5 - 4 in one circle, rarely 3, 2 or even 1 stamen. Gynoecium of 2-5 carpels, syncarpous or transitional to lysicarpous, mostly with free columns. Ovary superior, usually with numerous ovules in each nest, rarely with several ovules or only one. Fruits - boxes, nuts, rarely berries. The embryo is usually bent around a mealy perisperm.
Pollination is carried out mainly by insects. The flowers of paronychia and alsinaceae with free, spreading sepals and petals are not specialized for pollination by certain species; nectar and pollen in them are available to a wide variety of insects that carry out cross-pollination. In carnations, nectar, sometimes quite plentiful, is secreted by the expanded bases of the stamen filaments. Among the representatives of the family, good honey plants are known: common tar (Viscaria vulgaris), cuckoo adonis (Coro-naria flos-cuculi), lush carnation (Dianthus superbus). Smolevkovye flowers with soldered sepals are pollinated mainly by diurnal and nocturnal butterflies. With their long proboscises, butterflies get nectar from the bottom of the tubular cup, while they always touch the stamens, and the adhering pollen is transferred to other flowers. It is also important that the flowers of many carnations are painted in various shades of red, and butterflies, unlike many other insects, are able to perceive red.
The flowers of a number of carnations are protandric, in which the pollen ripens and spills out before the stigma of the same flower becomes capable of receiving it. Protandria is known from carnation grass (Dianthus deltoides), cuckoo chickweed, multi-flowered tar (Silene multiflora), green-colored tar (S. chlorantha) and many other cloves.
Not only protandria, but also such phenomena that are not uncommon among carnations, such as gynodietsia - female dioeciousness (some carnations, asterisks, tars) and trietia - three-domedness (certain types of soapwort - Saponaria - and tartar), practically eliminate self-pollination and contribute to more successful cross-pollination, not excluding, however, the possibility of geitonogamy.
Carnation flowers, pollinated by moths, are light in color and may be odorless or scented. The drooping tar (Silene nutans), growing in meadows, in well-lit places, opens white petals and emits a strong smell only in the evening, attracting night butterflies. Flowering lasts three nights, and in the maturation of stamens and stigmas, a certain sequence is observed that eliminates self-pollination: on the first night, the stamens of the outer circle ripen, on the second, the stamens of the inner circle, and only on the third night do the stigmas ripen. From crawling insects, which would only use nectar and pollen, but would not produce pollination, the drooping resin is protected by a sticky mass secreted by the stem at the base of the pedicels. As soon as flowering ends, the sticky substance ceases to stand out. Similarly, in many other cloves, such as common tar (Table 55), the stem is very sticky, which protects the flowers from uninvited guests - ants and other crawling insects. In the ovary of the soapwort (Saponaria officinalis), the cuckoo adonis, the drooping resin and some other clove butterflies, they lay eggs and pollinate. Caterpillars that appear soon feed on immature seeds, and then leave the flower, but there are quite a lot of seeds for reproduction in such plants.
Some members of the family, normally insect-pollinating plants, are able to switch to self-pollination if, due to bad weather or lack of insects, cross-pollination is not possible. In closed - cleistogamous flowers, which are found in certain species, for example, in lying bryozoan (Sagina procumbens), forest pseudostellaria (Pseudostellaria sylvatica), self-pollination always occurs.
Some paronychia (hernia - Herniaria, paronychia - Paronychia) and alsin (dival - Scleranthus) with flowers in which the petals are barely developed or completely absent are pollinated by the wind.
The fruits of the vast majority of cloves are multi-seeded boxes that open with cloves and are usually located at the top of the stem (Fig. 200). Ripe seeds do not spill out immediately, but in parts in different directions, when a gust of wind or the touch of an animal shakes the stem. The teeth of the pods of the carnation grass, drooping resin and some other carnations close in bad weather, and water cannot damage the seeds. In panicled kachima (Gypsophila paniculata), belonging to plants known as "tumbleweeds", the teeth of the boxes are always bent inward, and the gaps between them are very small, so the seeds can be dispersed from the boxes only with sufficiently strong gusts of wind. This usually occurs in autumn when the plant breaks away from the roots and its loose spherical bushes are blown over long distances by the wind. An original method of seed dispersal is known in Wilhelmsia physodes, which grows mainly in the Arctic regions of Siberia and North America. Its boxes with mature seeds usually break up into three swollen, membranous-leathery nests, which are dispersed by the wind (Fig. 200).
The seeds of some cloves are dispersed by ants; equipped with seedlings, the seeds of three-veined meringia (Moerhingia trinervia) are taken away by ants that eat the seedlings (Fig. 200). Often the seeds are carried by the wind, especially in species that produce seeds with a membranous border (Fig. 200).
One-seeded, indehiscent fruits - nuts are carried by wind or animals. Diwala nuts remain inside the growing cup (Fig. 200), it easily clings to the hair of animals, which spread the fruits. In addition, diwala nuts can be dispersed by wind. In the same way, the fruits of the genera Pteranthus (Pteranthus), Cometes (Cometes) and Sclerocephalus (Sclerocephalus) are spread. In these plants, after flowering, the twigs and leaves in each inflorescence grow, usually harden, and the whole plant with fruits becomes hard, prickly (Fig. 200). At the fruiting stage, the stems of paronychia and hernia become brittle, parts of plants with nuts are carried by the wind (Fig. 200); in paronychia, in addition to the calyx, rather large membranous bracts remain with the fruits, which facilitate the spread of fruits by the wind. The fruit of the berry blister (Cucubalus baccifer) is very peculiar - it is a shiny black dry berry (Fig. 200); birds may be involved in the distribution of such fruits.
The vast majority of cloves are annual or perennial herbaceous plants. However, in the family there are several medium-sized, 60 - 180 cm high, shrub species of the genus Schiedea, endemic to the Hawaiian Islands. Shrubs and semishrubs are found mainly among carnations growing in arid and mountainous regions of the temperate zone, in the tropics and subtropics.
For some perennial forms of cloves growing in the tundra, high in the mountains, in semi-deserts and deserts, a peculiar pillow form of growth is characteristic. The stem of such plants near the soil itself repeatedly branches, while numerous shoots depart in different directions, which, in turn, repeatedly branch. The whole plant takes the form of a hemisphere or pillow, often prickly. Very thorny cushions are formed by some species of prickly prickly (Acanthophyllum, pl. 56), growing mainly in South Transcaucasia, Iran, Afghanistan and Central Asia. The leaves of these pillow plants are turned into thorns. Pillows are loose or compact; both grow very slowly; per year, the shoots lengthen by only a few millimeters, and the age of individual specimens is calculated in the centuries. A dense pillow is better warmed by the sun's rays, its shoots are well protected from the drying effect of the wind, and its own microclimate is created inside the pillows. Observations on the cushion plant of the sandwort (Arenaria obtusifolia) in the mountains of Colorado (USA) showed that its leaf temperature was 12° higher than the ambient air temperature. Very large pillows, up to 2 m in diameter, are formed by aretium-shaped kachim (Gypsophila aretioides, Table 56), which grows on rocks at an altitude of up to 2000 m in southern Transcaucasia, mountainous Turkmenistan and northern Iran. The pillows of this plant are very hard, from a distance they look like a stone dressed with lichen. The mass of the pillow sometimes reaches 150 kg. With the extreme scarcity of vegetation, such pillows are used by local residents as fuel. Some of the highest flowering plants are species of the genus Pycnophyllum (Pycnophyllum) in the alpine zone of the Andes. Some of them form round, moss-like cushions up to 1 m in diameter, which can grow near the edge of eternal snow.
In alpine meadows, which are located high in the mountains, in close proximity to spaces covered with ice and snow, you can find many carnations - representatives of the genera Smolevka, gerbil, gerbil, Alsina (Alsine), Minuartia (Minuartia) and some others. Almost all alpine plants are perennials, barely rising above the ground. They have a highly developed root system, many have a dense pubescence or waxy coating and a thick cuticle on leaves and stems. Alpine meadows are extraordinarily beautiful, they are colored with a mass of various large and brightly colored flowers; bright coloring helps to attract pollinating insects, which are very rare in the mountains.
Among the cloves, especially annuals, there are many such plants that are malicious weeds of crops. These plants usually complete their development cycle quickly and produce a huge amount of seeds.
Sowing toriza (Spergula sativa), which previously inhabited only Eurasia, later became a cosmopolitan, infesting the crops of spring grain and tilled crops, as well as flax. One plant, blooming in July, soon produces up to 30,000 seeds, 10% of which are viable already in the current field season.
Common cockle - an annual with gray-felt pubescence of leaves and single large pink flowers, formerly widespread in the Mediterranean, penetrated to Australia and the Cape. Cuckoo is found in crops of cereals and flax, being a particularly dangerous weed of cereals, since its seeds contain 6.5% of the poisonous glycoside gitagin, or agrostemin, which acts on the heart, nervous system and destroys red blood cells. The admixture of cockle seeds in flour in an amount of 0.5% or more makes it bitter in taste and dangerous to health. However, cockle venom is harmless to sheep, birds and small rodents.
Well known as a ubiquitous and difficult to eradicate weed, mainly vegetable crops, medium stellate, or wood lice (Stellaria media). The life cycle of woodlice takes less than 40 days and gives 2-3 generations over the summer. The lower part of the stems, as well as autumn shoots, can overwinter and bloom soon after the snow melts. One plant produces up to 25,000 seeds, which remain viable for 8 years, and in some cases up to 25 years. This plant readily eats livestock, it is also used to feed chickens and domestic songbirds.
Most species of the family contain saponins - substances that, when shaken with water, give abundant foam. Saponins are present in all parts of the plant, but most of them are in the parenchymal cells of underground organs. Many cloves, for example, soapwort, dawn (Lychnis chalcedonica), prickly prickly (Acanthophyllum gypsophiloides), certain types of kachima, have long been popularly known under the name of "soap root" and have been used as a substitute for soap. The foam formed by saponins differs from soap foam - it does not contain alkali. The property of saponins to give abundant foam when shaken is manifested at a very low concentration, in some cases even at a dilution of 1: 10,000. Currently, this property of saponins is used in fire extinguishers, in the production of fizzy drinks, beer, halva. Saponins are used in perfumery in the manufacture of shampoos, in the textile industry for washing and bleaching woolen and silk fabrics, for which ordinary alkaline soap is inapplicable, in technology when enriching ores by the flotation method.
The use of cloves in medicine is also associated with the presence of saponins in these plants. For medicinal purposes, mainly two plants are used - medicinal soapwort and naked hernia (Herniaria glabra). However, saponins are far from harmless substances. It all depends on how they enter the human or animal body. Most saponins, entering the digestive tract through the mouth, do not have a toxic effect, but when directly introduced into the bloodstream, many saponins cause hemolysis - the destruction of red blood cells. One of the most valuable saponinos for the economy - prickly prickly prickly - for a long time served as a subject of wide export. As a result of many years of harvesting, its thickets in South Kazakhstan and in the republics of Central Asia have practically disappeared, so the development of the foundations for introducing this species into culture was an urgent problem. In Western Europe, other saponinoses are grown - holly pumpkin (Gypsophila acutifolia) and panicled pumpkin (G. paniculata).
The most remarkable and best known of the family is the extensive genus Dianthus, which includes about 300 species, widely represented in Europe, Asia, tropical and South Africa, some species are found in North America. The center of diversity of species of this genus is the Mediterranean. Many carnations are favored ornamental plants and are cultivated almost universally for their pleasant scent and usually bright corolla color, often double due to splitting of the stamens and turning them into petals.
The smell of clove flowers is reminiscent of the aroma of cloves - spices, which are dried buds of the clove tree (Syzygium aromaticum) from the myrtle family.
At the end of the 11th century Chinese annual carnation (Dianthus chinensis) penetrated European gardens, from which, by crossing with other species, many varieties were obtained with flowers of various colors, sometimes reaching 15 cm in diameter. Numerous double, semi-double, remontant varieties grown for cutting were obtained from perennial carnation garden, or Dutch (D. caryophyllus) comes from southern Europe. One of the least whimsical and winter-hardy carnations is the bearded carnation, which has compact inflorescences of various colors. The perennial carnation pinnate (D. plumarius) is widely cultivated, originally from Europe (from the Italian Alps to the Tatras). Almost stemless carnation (D. subacaulis) from Southwestern Europe, fragrant carnation (D. fragrans) from the Caucasus mountains, alpine carnation (D. alpinus) from the highlands of the Alps, blood-red carnation (D. cruentus) from Balkan Peninsula, D. monspessulanus carnation from the mountains of Central and Southern Europe. On the territory of the European part of the Soviet Union, carnation grass and carnation lush (D. superbus) are very common species. Carnation grass grows in dry, well-lit places, on sandy slopes and forest edges; its medium-sized, but graceful, bright, pink-red flowers are clearly visible among other plants. Lush carnation with a pale pink corolla, the petals of which are cut along the edges into narrow, linear lobes, can be found in sparse forests, on forest edges and glades, in meadows. Both of these carnations are successfully grown in gardens.
As ornamental plants, in addition to cloves, representatives of various genera of this family are used (Tables 55, 56, 57).
Annuals, biennials, or perennials, distributed over much of the Northern Hemisphere. Five-petalled flowers with tubular bracts and small, dense, pubescent foliage of many varieties of tares can decorate a rock garden, and tall S. dioica (S. dioecious) and S. fimbriata (S. fringed) enliven a corner of the garden in a natural style.
The species listed below are relatively cold-resistant perennials.
S. acaulis (S. stemless)
Evergreen perennial with very short peduncles, forming a continuous carpet of greenery and flowers. Flowers of light or dark pink color with a diameter of up to 1.5 cm bloom in early summer (the plant, however, does not always bloom profusely). The leaves collected in rosettes are bright green, linear, up to 1 cm long. The height and diameter of the plant are 5x15 cm.
Alpestris (S. alpine)
White flowers up to 2 cm in diameter with several large teeth on the petals bloom from early to mid-summer. A few light green leaves taper towards the top and reach a length of 5 cm. Herbaceous perennial, forming high air curtains from straight branched stems. The height and diameter of the plant is 30x20 cm.
Dioica (S. dioecious, Sandman red)
Reddish-pink flowers with bilobed petals 2 cm in diameter bloom in early summer and are collected in loose panicles. Egg-shaped bright green leaves sit on long petioles and reach a length of 5 cm. The height and diameter of the plant is 60x30 cm.
Elisabethae (S. Elizabeth)
A semi-evergreen perennial that gives a single peduncle in the second half of summer, on which, as a rule, only one lilac-pink flower with a diameter of 3 cm sits. The height and diameter of the plant is 15x20 cm.
Clove - one of the largest families in the order of clove. It contains approximately 80 genera and 2000 species. Carnations can be found on all continents of the globe, in a wide variety of habitats. Representatives of the family grow in the tundra, among the carnations there are many forest and meadow plants. They are also found in arid regions: in the steppes, semi-deserts and deserts. In the mountains, carnations rise to the alpine belt, and one of the species, creeping chickweed (Stellaria decumbens), was found in rocky crevices in the Himalayas at an altitude of 6000 m, much higher than other flowering plants of the highlands. Carnations are especially widely represented in the temperate regions of the northern hemisphere, with the largest number of genera and species concentrated in the Mediterranean, Western and Central Asia. In the composition of the floras of most parts of the Arctic, representatives of this family most often occupy the fifth place in terms of the number of species. Several genera of cloves are found in the southern hemisphere, in its temperate zone, as well as in the mountains of the tropical region. Even on ice-covered Antarctica, where there are practically no higher plants, on the Antarctic Peninsula of the mainland, along with the grass antarctic pike, one species of clove was found - the whale colobanthus (Colobanthus quitensis). A significant number of species and even genera of cloves are cosmopolitans. Such are the genera toriza (Spergula), moric (Spergularia), chickweed (Stellaria), chickweed (Cerastium), gerbil (Arenaria), etc. The whole variety of genera and species of cloves, in accordance with the family system proposed by F. Pax and K. Hoffman, is usually divided into three subfamilies: paronychia (Paronychioideae), alsin (Alsinoideae) and smolevkovye (Silenoideae); representatives of the subfamilies differ in the presence or absence of stipules and some structural features of the flowers. The leaves of the carnations are opposite, rarely alternate, simple, entire, often narrow - linear or linear-lanceolate, equipped with scaly stipules (most paronychia) or without stipules (alsinovye and resinous). The flowers are usually collected in dichasial inflorescences very characteristic of this family, either highly branched and loose (in Kachima species - Gypsophila), or more compact, umbellate or corymbose (bearded carnation - Dianthus barbatus); much less often single flowers (common cockle - Agrostemma githago). The flowers are actinomorphic, in most representatives 5-membered. Sepals 5, free, or almost free (paronychia and alsin) or fused into a tube (small, for example, tar-flapper - Silene vulgaris; often there are bracts close to the calyx. Petals are usually 5, always free; in some species, the petals are barely developed or Only in resinous petals have long nails (narrowed part of the petals) and limbs whole or divided into narrow lobes (widened part of the petals), and on the border between them sometimes there are petal-like outgrowths forming the so-called adnexal corolla or corolla, Stamens 10, arranged in two circles or 5-4 in one circle, rarely 3, 2 or even 1 stamen.Gynoecium of 2-5 carpels, syncarpous or transitional to lysicargic, mostly with free columns.Ovary superior, usually with numerous ovules in each nest, rarely with several ovules or only one.Fruits - boxes, nuts, rarely berries.The embryo is usually bent around mealy perisperm. Pollination is carried out mainly by insects. The flowers of paronychia and alsinaceae with free, spreading sepals and petals are not specialized for pollination by certain species; nectar and pollen in them are available to a wide variety of insects that carry out cross-pollination. In carnations, nectar, sometimes quite plentiful, is secreted by the expanded bases of the stamen filaments. Among the representatives of the family, good honey plants are known: common tar (Viscaria vulgaris), cuckoo adonis (Coronaria flos-cuculi), lush carnation (Dianthus superbus). Smolevkovye flowers with soldered sepals are pollinated mainly by diurnal and nocturnal butterflies. With their long proboscises, butterflies get nectar from the bottom of the tubular cup, while they always touch the stamens, and the adhering pollen is transferred to other flowers. It is also important that the flowers of many carnations are painted in various shades of red, and butterflies, unlike many other insects, are able to perceive red. The flowers of a number of carnations are protandric, in which the pollen ripens and spills out before the stigma of the same flower becomes capable of receiving it. Protandria is known from carnation grass (Dianthus deltoides), cuckoo's adonis, many-flowered tar (Silene multiflora), green tar (S. chlorantha) and many other cloves. Not only protandria, but also such phenomena that are not uncommon among carnations, such as gynodiecia - female dioeciousness (some carnations, asterisks, tars) and trietia - three-houseness (certain types of soapwort - Saponaria - and tartar), practically eliminate self-pollination and contribute to more successful cross-pollination, not excluding, however, the possibility of geitonogamy. Carnation flowers, pollinated by moths, are light in color and may be odorless or scented. Smolevka drooping (Silene nutans), growing in meadows, in places well lit by the sun, opens white petals and emits a strong smell only in the evening, attracting night butterflies. Flowering lasts three nights, and in the maturation of stamens and stigmas, a certain sequence is observed that eliminates self-pollination: on the first night, the stamens of the outer circle ripen, on the second, the stamens of the inner circle, and only on the third night do the stigmas ripen. From crawling insects, which would only use nectar and pollen, but would not produce pollination, the drooping resin is protected by a sticky mass secreted by the stem at the base of the pedicels. As soon as flowering ends, the sticky substance ceases to stand out. In the same way, in many other cloves, such as common tar, the stem is very sticky, which protects the flowers from uninvited guests - ants and other crawling insects. In the ovary of the soapwort (Saponaria officinalis), the cuckoo adonis, the drooping resin and some other clove butterflies, they lay eggs and pollinate. Caterpillars that appear soon feed on immature seeds, and then leave the flower, but there are quite a lot of seeds for reproduction in such plants. Some members of the family, normally insect-pollinating plants, are able to switch to self-pollination if, due to bad weather or lack of insects, cross-pollination is not possible. In closed - cleistogamous flowers, which are found in certain species, for example, in lying bryozoan (Sagina procumbens), forest pseudostellaria (Pseudostellaria sylvatica), self-pollination always occurs. Some paronychia (hernia - Hermana, paronychia - Paronychia) and alsin (divala - Scleranthus) with flowers in which the petals are barely developed or absent at all, are pollinated by the wind. The fruits of the vast majority of cloves are multi-seeded boxes that open with cloves and are usually located at the top of the stem. Ripe seeds do not spill out immediately, but in parts in different directions, when a gust of wind or the touch of an animal shakes the stem. The denticles of the pods of the clove grass, drooping tar and some other clove close in bad weather, and water cannot damage the seeds. In panicled kachima (Gypsophila paniculata), belonging to plants known as "tumbleweeds", the teeth of the boxes are always bent inward, and the gaps between them are very small, so the seeds can be dispersed from the boxes only with sufficiently strong gusts of wind. This usually occurs in autumn, when the plant breaks away from the root and its loose spherical bushes are carried by the wind and over long distances. An original method of seed dispersal is known in Wilhelmsia physodes, which grows mainly in the Arctic regions of Siberia and North America. Its boxes with mature seeds usually break up into three swollen membranous-leathery nests, which are dispersed by the wind. The seeds of some cloves are dispersed by ants; equipped with a seedling, the seeds of three-veined meringia (Moerhingia trinervia) are taken away by ants that eat the seedling. It is not uncommon for seeds to be carried by the wind, especially in species that produce seeds with a membranous border. Single-seeded, overlapping fruits - nuts are dispersed by wind or animals. Diwala nuts remain inside the growing cup, it easily clings to the hair of animals, which spread the fruits. In addition, diwala nuts can be dispersed by wind. In the same way, the fruits of the genera Pteranthus (Pteranthus), Cometes (Cometes) and Sclerocephalus (Sclerocephalus) are spread. In these plants, after flowering, the twigs and leaves in each inflorescence grow, usually harden, and the whole plant with fruits becomes hard, prickly. At the fruiting stage, the stems of paronychia and hernia become brittle, parts of plants with nuts are carried by the wind; in paronychia, in addition to the calyx, rather large membranous bracts remain with the fruits, which facilitate the spread of fruits by the wind. The fruit of the berry blister (Cucubalus baccifer) is very peculiar - it is a shiny black dry berry; birds may be involved in the distribution of such fruits. The vast majority of cloves are annual or perennial herbaceous plants. However, in the family there are several medium-sized, 60-180 cm high shrubs - species of the genus Shidea (Schiedea), endemic to the Hawaiian Islands. Shrubs and semishrubs are found mainly among carnations growing in arid and mountainous regions of the temperate zone, in the tropics and subtropics.
For some perennial forms of cloves growing in the tundra, high in the mountains, in semi-deserts and deserts, a peculiar pillow form of growth is characteristic. The stem of such plants near the soil itself repeatedly branches, while numerous shoots depart in different directions, which, in turn, repeatedly branch. The whole plant takes the form of a hemisphere or pillow, often prickly. Very prickly cushions form some species of prickly prickly (Acanthophyllum), growing mainly in South Transcaucasia, Iran, Afghanistan and Central Asia. The leaves of these pillow plants are turned into thorns. Pillows are loose or compact; both grow very slowly; per year, the shoots lengthen by only a few millimeters, and the age of individual specimens is calculated in the centuries. A dense pillow is better warmed by the sun's rays, its shoots are well protected from the drying effect of the wind, and its own microclimate is created inside the pillows. Observations on the cushion plant Arenaria obtusifolia in the mountains of Colorado (USA) showed that its leaf temperature was 12° higher than the ambient air temperature. Very large cushions, up to 2 m in diameter, are formed by the aretiform kachim (Gypsophila aretioides, Table 56), which grows on rocks at an altitude of up to 2000 m in South Transcaucasia, mountainous Turkmenistan and Northern Iran. The pillows of this plant are very hard, from a distance they look like a stone dressed with lichen. The mass of the pillow sometimes reaches 150 kg. With the extreme scarcity of vegetation, such pillows are used by local residents as fuel. Some of the highest flowering plants are species of the genus Pycnophyllum (Pycnophyllum) in the alpine zone of the Andes. Some of them form round, moss-like cushions up to 1 m in diameter, which can grow near the edge of eternal snow.
In alpine meadows, which are located high in the mountains, in close proximity to spaces covered with ice and snow, you can find many carnations - representatives of the genera Smolevka, gerbil, gerbil, Alsina (Alsine), Minuartia (Minuartia) and some others. Almost all alpine plants are perennials, barely rising above the ground. They have a highly developed root system, many have dense pubescence or waxy coating and a thick cuticle on leaves and stems. Alpine meadows are extraordinarily beautiful, they are colored with a mass of various large and brightly colored flowers; bright coloring helps to attract pollinating insects, which are very rare in the mountains.
Among the cloves, especially annuals, there are many such plants that are malicious weeds of crops. These plants usually complete their development cycle quickly and produce a huge amount of seeds.
Sowing toriza (Spergula sativa), which previously inhabited only Eurasia, later became cosmopolitan, infesting spring grain and row crops, as well as flax. One plant, blooming in July, soon produces up to 30,000 seeds, 10% of which are viable already in the current field season.
Common cockle - an annual with gray-felt pubescence of leaves and single large pink flowers, formerly widespread in the Mediterranean, penetrated to Australia and the Cape. Cuckoo is found in crops of cereals and flax, being a particularly dangerous weed of cereals, since its seeds contain 6.5% of the poisonous glycoside gitagin, or agrostemin, which acts on the heart, nervous system and destroys red blood cells. The admixture of cockle seeds in flour in an amount of 0.5% or more makes it bitter in taste and dangerous to health. However, cockle venom is harmless to sheep, birds and small rodents.
Well known as a ubiquitous and difficult to eradicate weed, mainly vegetable crops, medium stellaria, or wood lice (Stellaria media). The life cycle of woodlice takes less than 40 days and gives 2-3 generations over the summer. The lower part of the stems, as well as autumn shoots, can overwinter and bloom soon after the snow melts. One plant produces up to 25,000 seeds, which remain viable for 8 years, and in some cases up to 25 years. This plant readily eats livestock, it is also used to feed chickens and domestic songbirds.
Most species of the family contain saponins - substances that, when shaken with water, give abundant foam. Saponins are present in all parts of the plant, but most of them are in the parenchymal cells of underground organs. Many cloves, for example, soapwort, dawn (Lychnis chalcedonica), prickly prickly (Acanthophyllum gypsophiloides), certain types of kachima, have long been popularly known under the name of "soap root" and have been used as a substitute for soap. The foam formed by saponins differs from soap foam - it does not contain alkali. The property of saponins to give abundant foam when shaken is manifested at a very low concentration, in some cases even at a dilution of 1:10,000. Currently, this property of saponins is used in fire extinguishers, in the production of fizzy drinks, beer, halva. Saponins are used in perfumery in the manufacture of shampoos, in the textile industry for washing and bleaching woolen and silk fabrics, for which ordinary alkaline soap is inapplicable, in technology when enriching ores by the flotation method.
The use of cloves in medicine is also associated with the presence of saponins in these plants. For medicinal purposes, mainly two plants are used - medicinal soapwort and naked hernia (Herniaria glabra). However, saponins are far from harmless substances. It all depends on how they enter the human or animal body. Most saponins, entering the digestive tract through the mouth, do not have a toxic effect, but when directly introduced into the bloodstream, many saponins cause hemolysis - the destruction of red blood cells. One of the most valuable saponinos for the economy - prickly prickly prickly - for a long time served as a subject of wide export. As a result of many years of harvesting, its thickets in South Kazakhstan and in the republics of Central Asia have practically disappeared, so the development of the foundations for introducing this species into culture was an urgent problem. In Western Europe, other saponinoses are grown - holly pumpkin (Gypsophila acutifolia) and panicled pumpkin (G. paniculata).
The most remarkable and best known of the family is the vast genus Dianthus, which includes about 300 species, widely represented in Europe, Asia, tropical and South Africa, some species are found in North America. The center of diversity of species of this genus is the Mediterranean. Many carnations are favored ornamental plants and are cultivated almost universally for their pleasant scent and usually brightly colored corolla, often double due to splitting of the stamens and turning them into petals.
The smell of clove flowers is reminiscent of the aroma of cloves - spices, which are dried buds of the clove tree (Syzygium aromaticum) from the myrtle family.
At the end of the XVIII century. Chinese annual carnation (Dianthus chinensis) penetrated European gardens, from which, by crossing with other species, many varieties were obtained with flowers of various colors, sometimes reaching 15 cm in diameter. Numerous double, semi-double, remontant varieties grown for cutting were obtained from perennial carnation garden, or Dutch (D. caryophyllus) comes from southern Europe. One of the least whimsical and winter-hardy carnations is the bearded carnation, which has compact inflorescences of various colors. The perennial carnation pinnate (D. plumarius) is widely cultivated, originally from Europe (from the Italian Alps to the Tatras). Almost stemless carnation (D. subacaulis) from Southwestern Europe, fragrant carnation (D. fragrans) from the Caucasus mountains, alpine carnation (D. alpinus) from the highlands of the Alps, blood-red carnation (D. cruentus) from Balkan Peninsula, Montpellier carnation (D. monspessulanus) from the mountains of Central and Southern Europe. On the territory of the European part of the Soviet Union, carnation grass and carnation lush (D. superbus) are very common species. Carnation grass grows in dry, well-lit places, on sandy slopes and forest edges; its medium-sized, but graceful, bright, pink-red flowers are clearly visible among other plants. Lush carnation with a pale pink corolla, the petals of which are cut along the edges into narrow, linear lobes, can be found in sparse forests, on forest edges and glades, in meadows. Both of these carnations are successfully grown in gardens.
A. N. Ponomarev, E. I. DemyanovaDichogamy is a functional dichotomy. It is caused by the maturation and exposure of pollen and stigma in flowers at different times, as a result of which the latter act either in the male (anther) or in the female (stigma) phase. Dichogamy manifests itself in the form of protandry (with earlier maturation of pollen) or protogyny (with earlier stigma). The simultaneous maturation of pollen and stigmas in a flower is called homogamy.
In bluebells, the anthers open while still in the bud. Pollen surrounds the style with a muff, holding on to the hairs covering it. Empty and dried anthers are visible at the bottom of the corolla. The lobes of the stigma are still quite closed at this moment. About a day after the blooming of the flower, the stigma lobes diverge and it becomes possible to pollinate them with foreign pollen brought by insects. But at the end of flowering, autogamy is also possible, due to the fact that the stigma lobes, twisting spirally downwards, touch the receptive surface of the column, which has retained its own pollen.
Rice. 32. Protogyny at Kornut's plantain (Plantago cornutii):
1
- a flower in the bud stage; 2
- the appearance of a stigma; 3
- wilting of the stigma; 4
- the opening of the flower and the beginning of the nomination of the stamens with the stigma already withered; 5
- staminate stage of the flower.
In tares (Silene multiflora, S. chlorantha, etc.), autogamy is excluded. They flower and pollinate at night. The flowers bloom at 18-19 hours, and close in the morning. On the first evening, when the flower blooms, five stamens are exposed, on the second evening - the next five stamens, fading by morning, and finally, at night, on the third day, stigmas come forward. Thus, the anther and stigma phases are separated in the flowers of the resins, but geitonogamy is possible, since flowers in different developmental phases can occur on individuals.
Protandria is a very effective means of preventing self-pollination in Umbelliferae and Villusaceae.
In Umbelliferae, the protandry is strict, impeccable, and covers not only individual complex umbrellas, but the entire individual as a whole. In most species of umbelliferes, this is achieved due to the strict order in flowering of umbrellas of different orders and its complete consistency (synchronism) in umbrellas of a given order. As a result, each individual sequentially appears several times now in the anther, then in the stigma phase, and these phases are sharply demarcated and, as a rule, do not overlap each other. An example is the medium cut (Libanotis intermedia). This type of protandria (Libanotis phylum), with multiple alternations of staminate and stigma phases, is very common in Umbelliferae. Much less often they have another type of protandria (type Peucedanum), characterized by a single phase change in all simultaneously and consistently flowering umbrellas of an individual, regardless of which order they belong to. Such a protandry is known so far in Lyubimenko's boletus (Peucedanum lubimenkoanum).
A sharp protandry of the entire inflorescence, sequential and simultaneous change of phases in the flowering of inflorescences of different orders is also characteristic of some tufts: field barkwort (Knautia arvensis), pale yellow scabiosa (Scabiosa ochroleuca), etc. As a result, autogamy and geitonogamy are impossible in Umbelliferae and tufts , cross-pollination is inevitable.
Rice. 33. Protogyny in Gerard's rush (Juncus gerardii):
1
- flower in the stigma phase in the evening (protogyny); 2
- an open flower in the morning at the time of pollination.
Protogyny is well expressed in cruciferous, rose, barberry, honeysuckle, etc. In many cases, the difference in the time of maturation of the stigma and pollen is so insignificant that the presence of protogyny seems doubtful. Protogyny is more pronounced in wind-pollinated plants, and not only in bisexual, but also in monoecious and dioecious plants. This is the case with rushes, sedges, cereals, wormwoods, plantains (). In the rushes, the population of the species enters the stigma phase from the evening of the previous day, and the next morning it passes into the stamen phase, and the dispersion of pollen is limited to 2-3 hours (). In monoecious sedges, the stigma phase precedes the anther phase by 1-6 days, and in plantains by 4-6 days.
In such cases, the role of protogyny is that the early exposure of the stigmas represents a preparation for very rapid wind pollination during the short diurnal periods of pollen dispersal. The same function is performed by protogyny in some desert haze.
The most effective means of preventing self-pollination, of course, is self-incompatibility. It is expressed in the absence of pollen germination on the stigma or in the cessation of the growth of pollen tubes in the style during self-pollination. Self-incompatibility in angiosperms is more widespread than dioecy. Compared with the latter, it provides a higher seed productivity, since in this case each flower can produce seeds, while in dioecious plants only half of the flowers. In addition, self-incompatibility ensures maximum economy of female gametes, since the possibility of unsuccessful combinations of gametes is prevented before fertilization. Self-incompatibility is regulated by genetic mechanisms. Distinguish between homomorphic and heteromorphic self-incompatibility. Homomorphic self-incompatibility is the most common in nature: it has been recorded in approximately 10,000 angiosperm species from 78 families. Self-incompatibility in this case is not accompanied by morphological differences in the structure of the flower. If self-incompatibility is combined with heterostyle (heterostyle), then it is called heteromorphic.
Rice. 34. Heterostyly in primrose (Primula sp.):
1-4
- short-column form ( 1
- sectional view of a flower; 2
- a short style with a papillary surface of the stigma at low magnification; 3
4
- pollen grain); 5-8
- long-column form ( 5
- sectional view of a flower; 6
- a long style with a papillary surface of the stigma at low magnification; 7
- the same at high magnification; 8
- pollen grain).
The phenomenon of heterostyly consists in the fact that the species has two or three forms of flowers located on different individuals and differing in the length of the columns and the arrangement of stamens (dimorphic and trimorphic plants). In dimorphic primroses (Primula), the long-columnar form has flowers with a long column, the stigma of which is located in the corolla throat, and the stamens are lower, deep in the corolla. The short-columnar form has a reverse arrangement of columns and anthers (). Forms of heterostylic plants also differ in other ways. So, short-columnar flowers are characterized in comparison with long-columnar ones by larger pollen and smaller papillae of the stigma. Self-pollination in both forms, as well as cross-pollination between individuals of the same morphological type, produces an insignificant amount of seeds (self-incompatibility), while cross-pollination between plants with different column lengths is highly effective. Examples of dimorphic (distilled) species are lungwort, primrose, buckwheat, kermek, etc.
The trimorphic heterostyle loosestrife (Lythrum salicaria) has short, medium and long columnar flowers, distributed over different individuals. The stamens in the flower of each form correspond in length to the flower columns of the other two forms. Fertilization is most effective when the stigma of each flower form is pollinated with stamen pollen of the corresponding length from the other two flower forms, as was shown by C. Darwin (»
