The family has about 90 genera and at least 2,500 species, widely distributed in tropical, subtropical and temperate regions, mainly in Central and South America.
Representatives of the family are herbs, shrubs or small trees with alternate (sometimes opposite in the inflorescence area), simple leaves. The flowers are usually in axillary terminal inflorescences, bisexual, actinomorphic or, rarely, slightly zygomorphic. The calyx is usually 5-lobed or 5-partite, remaining, often enlarged during fruiting. The corolla is wheel-shaped to tubular, 5-lobed, rarely bilipped. There are usually 5 stamens or fewer in zygomorphic flowers (4-2); the anthers open longitudinally or through apical pores. The nectar disc is usually developed. The gynoecium usually consists of 2 carpels, rarely of 5 carpels, usually with an apical simple style with a bilobed stigma; the ovary is usually bilocular (sometimes false-3 or 5-locular) or rarely 5-locular, usually with numerous ovules. The fruit is a berry or a septic capsule; the fruit rarely decays (Fig. 218). Seeds with endosperm.
Nightshade flowers are pollinated by various insects, and in tropical countries also by birds and sometimes even mammals.
The nightshade family is divided into 2 subfamilies - the nolanaceae (Nolanoideae) and the nightshade family (Solanoideae). The first is often considered as an independent family of Nolanaceae. The Nolanaceae subfamily is relatively more primitive than the Solanaceae. It includes 2 close genera - Nolana (Nolana, about 75 species, distributed from Peru to Patagonia and the Galapagos Islands) and Alona (Alona, 5-6 species in Chile). These are herbs or small shrubs with alternate, entire, more or less succulent leaves, growing mainly along sea coasts. Their gynoecium consists of 5 carpels.
All other genera of the Solanaceae family are included in the extensive subfamily Solanaceae. It, in turn, is divided into 5 tribes. The most primitive tribe is considered to be the Nicandreae tribe, consisting of one monotypic genus Nicandra, living in Peru and Bolivia. This is an annual herb with coarse ribbed, rigidly pubescent stems, serrated or lobed leaves, single flowers, a 3-5-locular ovary and a rather multi-seeded berry, enclosed in a calyx that expands greatly during fruiting. Nikandra physalis (N. physalodes) has spread widely as a weed in many regions of our country in melon fields, vineyards, vegetable gardens and along roads, penetrating the Caucasus and oases Central Asia and on Far East. Occasionally it is bred for medicinal purposes and as an ornamental plant. Nicandra has an interesting adaptation against self-pollination. In the ephemeral flowers of this plant, the stigma withers within an hour after pollination and the style falls off.
The largest tribe of the nightshade family is the nightshade tribe proper (Solaneae), which includes several dozen genera. Their ovary is 2-locular, rarely multi-locular (for example, in a tomato). The tribe, in turn, is divided into several subtribes, of which the most primitive is the wolf subtribe. This includes trees, shrubs or grasses. The most famous and largest genus of wolfberry (Lycium) contains about 100 species of deciduous or evergreen erect or climbing, usually thorny shrubs, distributed in tropical, subtropical and partly temperate regions, mainly in South America. In our country there are 7 species in deserts, semi-deserts and steppes in the southeast of the European part, the Caucasus and Central Asia. Some species are bred for hedges and as ornamentals.
The subtribe of Derezoaceae also includes such a well-known genus as belladonna, or Atropa, consisting of 4 species distributed from Europe and the Mediterranean to India. The most famous belladonna, or belladonna (A. bella-donna, table 56), is a perennial herbaceous plant with a tall stem, flowers mostly with a brown-violet or dirty purple (sometimes yellow) corolla and multi-seeded black (sometimes yellow) shiny berry. Belladonna is a valuable medicinal plant introduced into culture. All parts of the plant contain alkaloids (atropine, hyoscyamine, etc.) and are poisonous. The attractive-looking belladonna berries are sometimes the cause of poisoning in children.
In the Solanaceae tribe, a separate subtribe also consists of the genera Scopolia (Scopolia) and Henbane (Hyoscyamus). Large perennial herbs with thickened rhizomes. Of the 6 species of scopolia, distributed from Central and Southern Europe to India, Tibet and Japan, only one species lives in our country growing wild - Carniolian scopolia (S. carniolica) in deciduous forests in the west of the European part and in the Caucasus. In all parts, but mainly in the rhizome and roots, it contains the alkaloids hyoscyamine and scopolamine. Other species - light yellow scopolia (S. lurida), found in the Himalayas, and Tangut scopolia (S. tangutica), growing in Tibet, also contain the alkaloid atropine. All 3 species are cultivated as valuable medicinal plants.
There are about 20 species in the genus henbane (Hyoscyamus), native to the Canary Islands, Europe, North Africa (to the central part), Western and Central Asia. In the USSR there are 8 species, of which highest value has black henbane (N. niger), which is harvested as a medicinal raw material in the European part of the USSR. Introduced into culture. Henbane contains alkaloids hyoscyamine, scopolamine, atropine and some others. The seeds of this henbane are similar in shape and color to poppy seeds and, when mixed with it, cause poisoning. Henbane pollination is cross-pollinated. The size of its dirty yellow, funnel-shaped corollas, with purple spots indicating the way to nectar, exactly corresponds to the size of larger bumblebees. Cross pollination is ensured by the fact that the stigma is located above the anthers.
The Solanaceae subtribe occupies a central position in the family tribe of the same name. Here we must first of all mention the rather large genus Physalis, about 100 species of which are widespread in tropical, subtropical and partly temperate regions, mainly in tropical America. Physalis are characterized by a very large, bubble-like, swollen red or orange calyx when fruiting. In the common physalis (P. alkekengi), the calyx of the fruit is especially large and bright, so this plant is very popular as an ornamental. The fruits of some species, such as common physalis and Peruvian physalis (P. peruviana), are quite edible and have a unique and pleasant taste.
The Solanaceae tribe also includes such a well-known cultivated plant as vegetable pepper (Capsicum annuum). In the botanical literature, the number of species of the genus Capsicum ranges from 20 to 50. Unfortunately, the taxonomy of this genus is in such a chaotic state that the number of its species cannot be determined. Species of capsicum (commonly called "vegetable pepper" in the literature, although not all of its species are vegetable plants) grow wild in Central and South America and the Galapagos Islands, as well as the Florida Peninsula. These are small shrubs, subshrubs or perennial herbs (usually used in cultivation as annuals) with multi-seeded red, orange or yellow, sometimes whitish or brownish fruits of various shapes. In tropical countries, several species of capsicum are widely cultivated, of which the common vegetable pepper, or paprika (C. annuum), is grown as an annual plant in countries with temperate and subtropical climates; in our country - in Ukraine, Moldova, the Caucasus and Central Asia. Its original wild perennial variety is distributed throughout the Florida peninsula, West Indies, Mexico, Central America and Colombia. The pungency of the fruits of the genus Capsicum depends on the phenolic volatile substance capsaicin. The capsaicin content is especially high in the fruits of Capsicum fruticosa (C. frutescens), the exact place of origin of which remains unknown. The seasoning obtained from the fruits of this species is known as Tabasco sauce. Capsicum shrub is cultivated in Central America, Mexico and the southern regions of the USA. The fruits of species of the genus Capsicum were widely used in America by the Aztecs, who daily flavored their food with finely crushed dry fruits. After his first voyage, Columbus brought capsicum from the West Indies. In Russia, the first mention of vegetable pepper is contained in the manuscript “Blessed Flower, or Herbalist” (1616).
But, of course, the most important for humans is the largest genus of nightshade (Solanum) in the family, numbering about 1,700 species, i.e., more than half of the species composition of the entire family. It is widespread in tropical, subtropical and temperate regions of both hemispheres, but mainly in South America. These are perennial, less often annual herbs, subshrubs with erect or climbing stems, and sometimes small trees. The fruit is a 2-locular multi-seeded berry. There are about 20 wild nightshade species in our country. Among them, bittersweet nightshade (S. dulcamara) is a climbing subshrub, mostly with purple flowers and bright red berries. It is found throughout almost the entire European part and in the south of Western Siberia. Another, even more common species in our country is black nightshade (S. nigrum) - an annual with white flowers and black (rarely green) berries, which grows as a weed in vegetable gardens and gardens and in weedy places. Both species contain the alkaloid solanine and are therefore poisonous and cause poisoning in humans and domestic animals. The nightshade genus includes a number of important cultivated plants. The first place among them is occupied by potatoes (from the German word Kartoffel). In culture, there are mainly 2 closely related species known - the Andean potato (S. andigena), which has long been cultivated in Colombia, Ecuador, Peru, Bolivia and Northwestern Argentina, and our ordinary tuberous potato (S. tuberosum), also called Chilean. Its homeland is Central Chile and adjacent islands (including the island of Chiloe). This species is very widespread in countries with temperate climates. The local population of the mountainous regions of South America also grows several other species. All types of potatoes belong to the tuberarium section of the nightshade genus, which, together with wild tuberous species, numbers about 200 species, growing mainly in South and Central America. Cultivated potato species are propagated by tubers (in breeding work, also by seeds). The introduction of potatoes into culture (first through the exploitation of wild thickets) began approximately 14 thousand years ago by the Indians of South America. Potatoes were first imported into Europe (Spain) around 1565, from where they spread to other countries. Potatoes first came to Russia in the 17th century, but the beginning of widespread potato cultivation was laid by a Senate decree in 1765 and the import of a batch of seed potatoes from abroad, sent throughout the country.
The process of pollination in potatoes is interesting. Its five stamens, folded together into a cone, fit tightly to the column protruding above them, the capitate stigma of which is slightly inclined downwards. When shaken, the anthers release a small amount of pollen. When visiting a flower, insects first touch the downward inclined stigma, and if they already had pollen from another flower, they pollinate it. But since only relatively few insects visit potato flowers, self-pollination usually occurs. Self-pollination occurs due to the fact that the stigma straightens and becomes exactly on the line along which the pollen falls.
Another very important cultivated representative of the nightshade genus is the eggplant, or badrijan (S. melongena). This is a perennial herbaceous plant with a tall stem, large leaves, purple flowers and more or less round, pear-shaped or cylindrical fruits. The fruits are yellow, with brown stripes, white, green or purple. Eggplant fruits are fried, stewed, pickled, and used to make eggplant caviar, sauté, etc. Eggplant grows wild in India and Burma. Eggplant was first introduced into culture in India, from where its culture spread to other countries, in particular to China. As early as 500 BC, a small-fruited form of eggplant was cultivated in China.
Among the cultivated nightshades, naranjilla (Solanum quitoense), the “golden fruit of the Andes,” should also be mentioned. Experts believe that naranjilla has a great future, although currently few people know about this wonderful plant outside of Colombia and Ecuador. This is an exceptionally tasty dessert fruit, which is also used for making jelly, jam and other purposes. Freshly squeezed juice from these fruits is used in Ecuador and Colombia to make “sorbete” - a green, foaming drink with an attractive sweet and sour taste of pineapple and strawberries. In Panama, Guatemala and Costa Rica, where the plant was introduced, the fresh juice is converted into a frozen concentrate. The plant is a shrub 1-2 m high, with pubescent leaves and round, yellow-orange fruits covered with easily removed white hairs. In favorable conditions, the plant bears fruit throughout the year.
Among the species of the extensive nightshade genus there are a number of other cultivated representatives.
The tomato, or tomato (Lycopersicon esculentum), cultivated in many varieties in all parts of the world, is very close to the nightshade genus and many botanists associate it with it. There are about 7 species in the tomato genus, living on the Pacific coast of South America (Colombia, Ecuador, Peru and Chile) and the Galapagos Islands. In the local Pahuatl language, this plant is called tomati, but when it was introduced in the 16th century. in Spain and Portugal they began to call it the “golden apple” (pomo d’oro - hence “tomato”).
The extensive nightshade tribe, but as a separate subtribe, includes the South American genus Cyphomandra and the Mediterranean-Asian genus Mandragora. There are about 6 species in the genus Mandrake, distributed from the Iberian Peninsula to the Eastern Himalayas and Tibet. Most species of mandrake are perennial herbs, almost always stemless, with very large leaves in a rosette, reaching a diameter of 1-2 m or more. The fleshy, starch-rich roots of mandrakes have a peculiar branching pattern: sometimes the root produces two vertical branches and somewhat resembles a human figure. Because of this feature, the mandrake has been covered in legends since ancient times, attributing to it magical power. In addition, it contains the alkaloid hyoscyamine and in the Middle Ages was considered one of the most valuable medicinal potions.
In Western Kopetdag, in the Turkmen SSR, a new species was discovered - the Turkmen mandrake (M. turcomanica). Here, the growing season of this plant begins in autumn, with the beginning of the rainy season, when it develops a rosette of large leaves. Flowering in conditions warm winters Western Kopetdag occurs in early November and continues until mid-April. Flowers are pollinated by various insects. Fruit ripening is observed from May to the end of June. With the onset of a hot, dry period, the plant seems to freeze and shed its dried leaves. The ripe fruits of the Turkmen mandrake are large (up to 5-6 cm in diameter), orange, aromatic and quite edible.
Unlike the stemless Mediterranean species of mandrake, the Himalayan-Tibetan stem mandrake (M. caulescens) has a developed stem and smaller leaves. At the same time, its closest related species, the Tibetan mandrake (M. tibetica), is a small, densely rosette plant. As one can assume, in this case, adaptive evolution followed the path of fixation of the juvenile phase of the vegetative sphere.
Next in the system of the Solanaceae subfamily comes the small tribe Datura (Datureae), characterized by the fact that as a result of the development of two false septa separating each of the two placentas of the initially two-locular ovary, the ovary becomes, as it were, four-locular. The fruit of Datura is a capsule or berry. Most well-known representative This tribe is the genus Datura, which numbers about 10 species of perennial or annual herbs, living in tropical and warm-temperate countries, mainly in tropical America. The flowers of Datura species are large, with a white funnel-shaped corolla from 6 to 20 cm or more in length. All of them are distinguished by large, long-tubular flowers with an intoxicating smell and are pollinated mainly by long-proboscis moths, which fly from everywhere to the smell of dope, neglecting other plants. In the process of evolution, the elongation of the corolla tube in many representatives of the genus paralleled the increase in the length of the proboscis of some hawkmoth butterflies (Sphingidae). As a result, species of Datura have emerged that can only be pollinated by certain species of hawk moths, becoming completely dependent on them. Datura fruits are original prickly capsules that open with four doors. These poisonous plants contain a number of alkaloids and are used in the pharmacopoeia, and were known to the ancient Peruvians as anesthetics. In some places, Datura species are cultivated as ornamental plants. In our country, the most widely known annual datura is the common or stinking dope (D. stramonium), which is found as a ruderal plant in wastelands, vegetable gardens, near hedges and buildings throughout almost the entire European part of the USSR. It is cultivated as a medicinal plant in the south of Ukraine and in the Krasnodar region. All parts of the plant contain poisonous alkaloids (hyoscyamine, scopolamine, atropine, etc.). In the southern regions of the USSR, harmless datura (D. innoxia) native to America and Indian datura (D. metel) native to Southwestern China are cultivated, the unripe fruits of which contain scopolamine.
The South American genus Brugmansia (Table 56) is very close to Datura and is often associated with it, consisting of 5 species of shrubs or small trees with very large leathery leaves up to 30 cm long. Huge bright tubular flowers hanging on long stalks open in the morning, and in the afternoon, in the heat, they slam shut again, which Drosophila species take full advantage of. The flies collect in the corolla immediately after it opens. Despite the fact that fruit flies are located in the flower near the anthers, they apparently do not take part in pollination. After several days, the corolla of a flower inhabited by flies falls off, and along with it, grown-up larvae of fruit flies fall to the ground, continuing to feed on the pollen remaining in the corolla. The fruit flies are probably finishing their life cycle together with the flower of the individual that sheltered them. G. Carson, who observed certain species of fruit flies settling in the flowers of Brugmansia white (B. candida), believes that these types of fruit flies can only exist in cohabitation with Brugmansia. Flowers are pollinated by larger insects that can penetrate the flower from bottom to top. The elongated, soft or somewhat woody, indehiscent fruits of Brugmansia contain large wedge-shaped seeds. “Tree-like dope” is widely known as an ornamental plant, which is a hybrid between snow-white Brugmansia and woody Brugmansia (B. arborea, table 56).
Among other representatives of the tribe, the extremely interesting genus Solandra should be noted, about 10 species of which are common in tropical America. Most solandras are long and thick-stemmed vines that live in tropical mountain forests and twine around large trees. Climbing up their trunks to a great height, the liana spreads simple, entire, leathery leaves at the top of the tree crown and exposes the sun to large, beautiful, slightly zygomorphic bell-shaped flowers on thick pedicels. The polyspermous berries of plants, freely covered by an overgrown bright calyx, are clearly visible from above on the surface of the crown of the host tree and therefore are readily eaten by birds, which carry their kidney-shaped small seeds over long distances.
Some individuals of Solandra in old age almost or completely lose contact with the soil, continuing to function as semi-epiphytes or as true epiphytes. The most beautifully flowering species of the genus are cultivated in tropical gardens and greenhouses as ornamental ones. Particularly popular is solandra grandiflora (S. grandiflora), which weaves a living carpet around fences and walls of houses.
It is interesting that pollination of some representatives of the Datura tribe is carried out not only by various insects, but also by bats. This has been traced to the genus Trianaea, 3 species of which are common in the mountain forests of the Northern Andes. These plants have rather large flowers located at the ends of long hanging branches. and secrete abundant nectar. The flowers open in the evening, emitting a sharp bad smell, which attracts bats.
The tribe of Cestreae was named after the most prominent genus Cestrum (Table 56), which unites about 150 species living in tropical and subtropical regions of America. These are shrubs or small trees with entire, mostly narrow, often pubescent leaves. Their flowers, collected in apical or axillary inflorescences with a small calyx and a long funnel-shaped or tubular corolla, open mainly at night, emitting a strong odor. Particularly distinguished by this is the nocturnal cestrum (C. nocturnum), called “night jasmine” in its homeland. This shrub, widely grown in tropical gardens, produces a huge number of small greenish-white to cream flowers, which at night emit a very pleasant and strong aroma, attracting nocturnal pollinating insects. L. Overland (1960), studying the mechanism of opening and closing of “night jasmine” flowers, came to the conclusion that the substances responsible for their strong night smell are localized in the parenchyma cells of the petal apex. The occurrence of smell is not the result of the onset of darkness, as one might assume, but a consequence of the internal rhythm of the plant, independent of external conditions. The opening and closing of young flowers during the day is synchronized with the cycle of the appearance and disappearance of the smell: flowers that are wide open (at night) smell, closed flowers (during the day) are not. As the flower ages, the cyclicity gradually breaks down; very old, already fertilized flowers do not smell and do not close. Some species of cestrum, such as C. campestre, are pollinated by hummingbirds. Cestrum fruits are oblong berries.
The genus tobacco, or Nicotiana, also belongs to this tribe, numbering 66 species, of which 45 live in extratropical parts of America, and 21 species are confined to Australia and Polynesia.
These are mainly annual herbs, occasionally perennial herbaceous plants, but shrubs are also found. Flowers with a large tubular funnel-shaped or bell-shaped corolla. Hummingbirds help pollinate tobacco in America. The fruit is an ovoid, 2-4-leaf capsule filled with numerous small reticulated punctate seeds. Plants with a strong unpleasant odor. Many species of the genus contain nicotine and other toxic alkaloids. In all countries where wild tobacco grows, cases of poisoning of their leaves and young shoots of domestic animals have been repeatedly observed.
In cultivation, the most famous are real tobacco (N. tabacum) and shag tobacco (N. rustica). Shag leaves are used not only for smoking, but they are also the main raw material for obtaining citric acid, nicotine and nicotinic medications - nicotinic acid (vitamin PP), nicotine sulfate, which is effectively used to control agricultural pests. Tobacco was grown in America long before its discovery by Europeans. Tobacco was brought to Europe at the end of the 15th and first half of the 16th centuries. and was first grown as an ornamental and medicinal plant.
Close to tobacco is the genus Petunia, represented by about 30 South American species; some of them are widely cultivated as ornamental plants. Our most famous plant is the hybrid petunia (P. hybrida).
The genus Markea, which belongs to the Cesteraceae, is extremely unique, 18 species of which live primarily in the forests of tropical America - from the Amazon River basin to Mexico. Some types of markea are vines with long graceful stems and simple, entire leaves, collected in bunches at the ends of the branches. Twisting around the support, the stems of the marquea creep higher and higher through the trees, forming terminal inflorescences with velvety short-tubular actinomorphic, often purple-green flowers in the upper part of the crown of the host tree. S. Vogel in 1958 observed that the flowers of some species of markea, such as markea dressleri (M. dressleri), are closed during the day and open only at night, attracting with their smell local forest rats, which are the main pollinators of these plants . Other species of the genus lead an epiphytic lifestyle, which is especially typical for curly marchea (M. ulei). This small shrub with inconspicuous, inconspicuous flowers is found in dense mature forests from Panama to Peru. Its individuals are able to settle both in the shady crown of a tree and at the top of the crown, under the open scorching sun. Thin aerial roots of epiphytic plants, covered with a loose shell, as well as water-permeable young stems and leaves intensively absorb moisture, oxygen, carbon dioxide, and even minerals, which enter the air along with tiny soil particles. Epiphytes develop intensively during the rainy seasons, accumulating water in their loose tissues, which they then use during the dry season. From time immemorial, these unique vessels of moisture and nutrients have been used by ants, arranging long-term shelters inside the stems of some types of markea. Symbiosis with ants is also beneficial for shrubs, since ants not only protect their living home from insect pests and other external enemies, but also bring nutrients for the plant to the substrate along with excrement.
The most advanced tribe of the family is the Salpiglossideae tribe. Plants belonging to it often have zygomorphic flowers with 2-4 fertile stamens and septicidal dehiscent capsule-type fruits with a lignified outer epidermis.
One of the notable genera of the tribe is Schizanthus, an endemic Chilean genus of about 10 species. It differs from other Solanaceae by its clearly defined two-lipped and extremely zygomorphic corolla. Of the 5 stamens, only 2 are fertile, 2 are transformed into staminodes, and the fifth is very reduced. The mechanism of flower pollination is interesting: two fertile stamens are located inside the lip formed by the two lower lobes of the corolla. When a bee or butterfly lands on this lip, the anthers explode vigorously, shooting pollen into the air. Some of the pollen grains usually land on an insect, which carries them to another flower. Some species of this genus have long been cultivated as ornamental plants. Particularly popular is pinnatus schisanthus (S. pinnatus, Fig. 219), a perennial glandular pubescent plant with lacy leaves and numerous variegated flowers, called the “butterfly flower” or “poor man’s orchid.” A number of forms have been developed in culture, differing in the color of the petals.
Another noteworthy genus of salpiglossia is Brunfelsia, which consists of 40 species native to tropical America. It is widespread in the tropical zone ornamental trees or shrubs that bloom during the rainy season and are therefore called “rain trees.” Brunfelsia leaves are entire-edged, leathery, often located at the ends of the branches, sometimes collected in bunches. The flowers are slightly zygomorphic, blue, purple or white, with 4 fertile stamens. With age, the flowers change color, like borage. The fruits of most species are juicy or leathery berries, surrounded by a slightly expanded calyx and containing large prismatic seeds with a fleshy endosperm. Some of the species are very decorative.
The genus Salpiglossis, which gave the tribe its name, lives in South America in 5 species. Salpiglossis notched (S. sinuata) is found as an ornamental plant in culture.
Life of plants: in 6 volumes. - M.: Enlightenment. Edited by A. L. Takhtadzhyan, editor-in-chief, corresponding member. USSR Academy of Sciences, prof. A.A. Fedorov. 1974 .
Elena Verkhovtseva
A series of educational events in preparatory group: “From the carriage to the rocket. From the distant past to the present."
Ulyanovsk
Educational event in the preparatory group on the topic: “Along the seas, along the waves.”
Program content.
1. Deepen children’s understanding of water transport (distinguish it by purpose, group it according to common characteristics). Reveal the importance of water transport in human life.
2. Introduce the first shuttle ships and other means of transportation on water (inflatable skins, rafts, logs, boats, etc.). Trace the history of shipbuilding: from small boats, one-trees, to modern motor ships and nuclear powered ships, submarines.
3. Arouse educational interest in the history of navigation. Introduce children to pirates and Vikings, who in ancient times were engaged in robbing merchant ships. Introduce children to the symbolism that existed in ancient times on pirate ships and Viking ships.
4. Consolidate existing knowledge about sea and river transport as a means of transportation, transportation of goods and use for other purposes.
5. Activate children's vocabulary: dugout boat, longship, sails, skerries, schooner, galleys, boarding, pirates - corsairs, privateers, privateer ship, Vikings, drakar, sailors, steamship, motor ship, nuclear icebreaker, tanker, Submarine etc.
Preliminary work.
Conversations on the topics: “Transport”, “Peter I - the founder of the Russian fleet”.
"Who to be?" (marine professions, “Gentlemen of Fortune.”
Conversation-game: “Who would you take with you on a ship?”, “Things and objects needed on a voyage.”
Acquaintance with the history of the legendary cruiser "Aurora". Learning the song “Cruiser Aurora” by V. Ya. Shainsky.
Examination of illustrations of ancient and modern modes of transport.
Excursion to the river port: acquaintance with the work of the river port, observation of boats, boats, cargo barges, passenger ships.
Reading of the poem: M. Yu. Lermontov “Sailboat”, M. V. Isakovsky “Go beyond the seas and oceans”.
Reading excerpts from books about sea travelers. Watch a video on this topic.
Learning songs about sailors with children.
Learning poems with children on a marine theme.
Reading the story “Flies high, but sees deep.”
Examination of I. E. Repin’s painting “Barge Haulers on the Volga.”
Listening to the Russian folk song "Volga".
Learning the song “Once upon a time there lived a brave captain” (I. O. Dunaevsky).
Proverbs and sayings about the Motherland, work, friendship.
Guessing riddles on the topic: “Water transport.”
Learning poems about ships.
Board game (puzzles) “Sailboat”.
Viewing the exhibits of the group museum.
Manual labor: "Steamboats".
Application on the theme “Across the seas, along the waves.”
Drawing: “Sailboats on the Volga”, “Transport of the future”.
Making crafts from waste material(traffic jams) “Water transport.”
Album design with illustrations, children's works, photographs, songs, poems on the theme “Blue Sea”.
Learning the sailor-themed dance “Apple”.
Games: “School of Sailors”, “Question and Answer”, “Connoisseurs”.
Outdoor game “The sea is agitated once.”
Travel game: “Submariners”.
Didactic game “Learn a profession.”
Production of attributes for the role-playing game (marine binoculars, caps). Role-playing game “Journey along the Volga”.
Interaction with family:
Cooking the dish “Navy-style pasta” with my mother.
Involving parents in the production of attributes for the role-playing game “We are walking along the Volga.”
Involving parents in the design of the illustrated album “Blue Sea”, “Our Army is Strong”.
Joint design of a plot painting on the theme “Sailboats on the Volga”.
Involving parents in the design of a mini-museum in a group (constructing boats, sailboats, frigates from plastic, plywood).
Involving parents in the production of attributes for the role-playing game “Journey along the Volga” (steering wheel, binoculars, telescope, compass).
Viewing by parents and children of feature films: “Pirates of the Caribbean”, “Children of Captain Grant” (the song “Once Upon a Brave Captain”) and cartoons: “Peter Pan”, “Treasure Island”, “Boat” (the song “Chunga-Changa”, "Captain Vrungel".
Material.
The book “The World of Ships”, “Children’s Encyclopedia”, layouts, illustrations, a geographical map, a board game with “Treasure Island” chips, a toy book with “Pirates” stickers, a cocked hat, a spyglass, a bottle with a map-plan indicating the location hidden treasures.
1. Educator: “Guys, today we will go on a trip in a time machine again. But the journey will not be easy, and therefore I suggest putting on life jackets and taking life preservers. Where do you think we will go? (Children's answers.) Tell me, what else needs to be taken on the road? (Game “Pack a Backpack.”)
Go boldly on your journey,
Mind, don’t forget your ingenuity.
Many difficulties will have to be overcome,
My friends and I need to be on time everywhere.
We are going back to the past of water transport. Here is our time machine, take your seats! (Shows a clock depicting the past and present of water transport.) What do we need to do to get to the past? (Turn the clock back.) Let's go! (The teacher or child moves the clock hand. Fast music plays.)
2. Guys, imagine that you and I find ourselves on a desert island. There is no food on the island, no bananas, no coconuts, no other fruits, in general, nothing except trees and flowers. There is another island nearby, it has everything. The sea separates these islands. If you and I don’t find a way to get to another island, we’ll die of hunger. What should I do? We have to cross the sea, but we don’t know how to swim. (And we are wearing life jackets and buoys) Good, but you can’t swim far in them, in case they burst or fall down, and the second island is very far away. What do we do? (Children's options.) So, you suggest using fallen trees. But it’s dangerous to swim holding on to the trees, there are sharks in the water. (You need to lie on a tree.) The tree, if there is only one, will turn over. (We need to tie the trees.) With what? There are no ropes. (You need to weave braids from the grass and tie them together, you’ll get a raft.) Well done!
3. Ancient man also thought about how to cross the sea. First I tried to inflate the skin, then I decided to float on a log, then I cut down several trees, cleared them of branches, tied them up - it turned out to be a raft. I used one of the branches as an oar. (Show illustration.) People have long noticed that tree trunks do not sink in water. Having tied them together and armed with a long pole, they ventured on their first voyages along the coast. Guys, do you think it’s possible to go on a long journey on a raft? (No, why? (Children's answers.) Because large waves can sink a raft or wash a traveler out to sea. The raft was a clumsy and heavy structure, but it was quite suitable for transporting large loads, especially if the voyage took place downstream. In deep places where the pole did not reach the bottom, people learned to steer the raft using a rowing board (perhaps this idea was suggested by observations of waterfowl). However, the raft could not satisfy all the needs of a person, who very often felt the need for a small, light and maneuverable floating device. The raft is considered one of the most primitive means of transportation on water.
There are several reasons that prompted a person to master water element. Ancient people often moved from one place to another and had to carry their belongings on themselves during their wanderings. Trying to make this difficult work easier, they began to think about means of transportation and, above all, learned to use the power of water to their advantage. In addition, in places that were located on the shores of seas or large rivers rich in fish, swimming equipment was necessary for fishing. Such a means was a wooden dugout boat.
First they felled the tree, then they supplied it with shovels.
Then they hollowed out his insides and let him walk along the river.
Its prototype was also a log. It was stronger and more reliable than a raft. It is not so dangerous to go out into the open sea. Whether a person burns the core out of a tree or chops it out with an ax, the boat is ready. Or rather shuttle. Then the oars appeared. (Show illustrations.) This boat moved faster than the raft, but, unfortunately, it often capsized.
4. U different nations The shapes of the boats and oars were different, and the materials from which they were made were also different. The first large ships appeared in Egypt and China. Egyptian ships were narrow, graceful, made of reeds with a papyrus sail, which helped them sail with a fair wind. When they sailed against the wind, they used oars. Then the Arabs invented their dhow, and the Chinese invented their junk. (Show illustrations.)
Man's fear of water comes from ancient times. Having overcome it and made the river its companion, humanity actively used water flows: streams, rivers and lakes, and later seas and oceans to meet the needs of civilizations. Floating on logs and waterskins, hollowing out a canoe from a tree trunk, placing a sail over a boat, people used rivers not only for fishing. The rivers were roads; in the summer, boats slid along their surface, and in the winter, sleighs with luggage rode.
5. The highest achievements of sailing shipbuilding were clippers and galleys. Galleys - large sailing and rowing ships - were built starting from the 8th century AD, and until the 18th century they were used in the fleets of almost all European countries. (Show illustrations.) Since the 14th century, galleys have been equipped with cannons. And yet, despite the combination of sails and oars, the galleys were rather clumsy, and their speed did not exceed seven knots (about ten kilometers per hour). Clippers are fast, lightweight, with powerful sailing rigs and excellent seaworthiness. (Show illustrations.) Clipper ships transported “noble cargo” - tea, spices, expensive oriental fabrics.
Long journeys across the seas and oceans became possible thanks to the creation of more advanced and reliable sailing ships, navigational instruments, instruments and maps that made it possible to navigate the open sea far from the coast. Geographical discoveries caused not only an unprecedented development of trade with distant countries, but also many wars between European states. Therefore, even merchant ships of that time had full military weapons - artillery and equipment for boarding combat.
In the 15th-17th centuries, the shape of the ships' hulls changed noticeably: they became more spacious, with a high bow and stern. It was simply impossible to move such a ship with the help of oars - it could only float thanks to its numerous sails. However, sailing-rowing galleys and galleasses have changed little - the shape of the hull was inherited from their ancient “ancestors”. In the 16th and 17th centuries, galleons were part of the fleets of England, France and Spain. These warships, with a displacement of up to 1000 tons and a length of up to 50 meters, were armed with 50-80 cannons and represented a formidable force. The living quarters on the galleon were located in a high aft superstructure with up to seven decks. The seaworthiness of the galleons was not the best - due to the high sides and bulky aft rooms.
6. Guys, look, someone left their things here. Who do you think? (Pirates.) Guys, who are the pirates? (People who went out on ships on the high seas to rob other ships.) What films about pirates do you know and have seen? (Children's answers.) Sea robbers arose at the very time when merchant ships appeared on the sea routes. Bloodthirsty adventurers and daredevils terrified peaceful merchants and travelers. Who knows what they were often called? (Gentlemen of fortune.) And there were pirates who attacked enemy ships with the permission of the king - however, this only happened during the war. Such pirates were called corsairs or privateers. And their ship was called a privateer ship. There were even women among the pirates. (Looking at illustrations and models.) The pirate crew often consisted of sailors who did not like the hard naval labor and strict discipline: in search happy life they fled to pirate ships. But among them there were also prisoners from captured ships who became pirates under duress.
Guys, you and I know that all ships have flags. Did the pirates have their own flag? What did he look like and what was his name? (Look at the illustration.) The flags were different, but flags with a skull and crossbones were more common. This flag was considered a pirate flag throughout the world and was known as the Jolly Roger. The flag was supposed to intimidate the victims and show what fate awaited them. For example, a skull with crossbones and an hourglass underneath meant that the victim's time had expired and the hour of his death had come. (The teacher shows the children pirate flags, and the children guess what they mean.)
Pirates were almost always armed to the teeth. What pirate equipment do you know? (Children's answers.) They could not do without a cutlass, a hook and an ax, as well as a pistol and a dagger. One deft throw of the boarding ax - and the cable (as thick as an arm) is cut, and this is enough for the enemy’s sails to fall. Pirate ships were small and very maneuverable. (Looking at illustrations.) The pirates tried to get closer to their victim as unnoticed as possible. Then they quickly threw grappling hooks onto the enemy ship. As soon as the ships approached, the pirates flew on board the captured ship with wild screams, and a terrible fight ensued: knives and pistols were used. What other weapons were on the pirate ship? (Cannons, rifles, grenades.) The Caribbean Sea, the Mediterranean Sea, the coastal waters of North Africa and Arabia, as well as the South China Sea, where pirates are still found, were considered the most dangerous for civilian ships.
Guys, do you know how pirates lived on a ship? (The teacher summarizes the children’s answers.) Life on board the ship, even for a pirate, was not easy. In a storm or any bad weather, one had to climb up to take care of the rigging (this is the equipment of the ship, install or remove the sails. This is sometimes even life-threatening. After a storm or battle, much on board needed repairs. It was necessary to patch the sails, strengthen the cables, repair masts and deck boards. From bloodthirsty robbers, pirates turned into ship carpenters. And during calm periods, the pirates suffered from idleness and quarreled among themselves. Life in difficult conditions soon brought the pirates into an unsightly state. They became dirty and smelled bad. During the voyage they had to sleep in a damp hold full of rats, they had no vegetables or fruits, and due to a lack of vitamins, the pirates suffered from scurvy. Having neither a doctor nor medicine on board, many of them could only replace a severed leg with a wooden prosthesis or put an eye patch on the missing eye.Pirates are bandits who travel on water.Although most pirates targeted and destroyed ships, some also attacked coastal cities.
Oh, guys, look, a bottle has sailed to our shore. Let's see what's there! (They find a “treasure map” - a plan of a group room, with the location of the “treasure” marked. The children, together with the teacher, follow the map and find a chest containing two games about pirates: “Pirates” - a toy book with stickers and “Island treasures" - a game with chips. The teacher offers to play them later, when they return back to the present time)
7. Guys, who else terrified civilian and merchant ships, as well as residents of cities and villages? (Vikings.) What do you know about the Vikings? (Children's answers.) They first appeared in Denmark, Sweden and Norway. (Display on a geographical map or globe.) When their drakarov ships approached foreign shores, the local residents trembled with fear. Most of the warships were brightly painted. Carved dragon heads, sometimes gilded, adorned the bows of ships. The same decoration could be on the stern, and in some cases there was a writhing tail of a dragon. This is where the name of the Viking ship came from. The drakars were long and narrow. They were very convenient for entering and navigating narrow river mouths. They were also stable enough to withstand sea waves. Viking ships moved with the help of sails and oars. The simple, square-shaped sail, made from rough canvas, was often painted with stripes and checkered patterns. The mast could be shortened and even removed altogether. With the help of skillful devices, the captain could steer the ship against the wind. The ships were controlled by a blade-shaped rudder mounted on the stern on the starboard side. The military leader stood bravely at the bow of the ship. What were the sides of the ships covered with? (Shields.) The sight of these fast ships with curved snake-like prows and warriors on board brandishing swords and axes was truly frightening. On the shallows near the shore, the warriors quickly jumped to the ground and easily pulled their ships ashore.
The Vikings often raided cities, villages, churches and monasteries. They took everything of value, set fire to houses and injured or killed anyone who stood in their way. When faced with an opposing army, the Vikings lined up tightly in ranks, shielded by a wall of shields. Guys, what weapons did the Vikings have? (Children's answers.) They attacked the enemy with arrows and spears. Then hand-to-hand combat with swords and axes began. One of the Viking soldier's favorite weapons was the battle axe. This ax frightened enemies with its sharp, deadly blade.
8. The development of water transport was a very important moment, which had a strong impact on various aspects of people’s lives. The voyages of merchants and military expeditions broadened the horizons of mankind and introduced them to new types of economic activity, tools, and food crops. Sea voyages and great geographical discoveries dramatically changed people's ideas about the world in which they lived and significantly contributed to the progressive development of mankind. Expanding connections between different territories and continents had and negative sides- they were the cause of the spread of epidemics of very serious infectious diseases.
Guys, who was the founder of the Russian fleet? (Peter I.)
This small sailing ship was called a boat. (Show illustration.) Russian soldiers boarded their fast boats, set sail and set off. Then the big ships appeared. They could have 2, 3 or 4 masts and many sails. Sailing ships that plied the seas in the past have very beautiful names. Depending on the number of sails and their location, they are distinguished: schooners, caravels, brigantines, frigates. (Looking at illustrations and models of ships.)
Frigates were the most advanced ships of the 17th century. These light, fast and maneuverable ships could carry enough cargo and still maintain mobility. Therefore, frigates were valued by the military, merchants, and sea pirates. The streamlined hull, the absence of high deck superstructures, two batteries of guns - one below deck and the other open - all these advantages gave the ship a long life. Look at the photo of frigates, these are magnificent snow-white beauties. (Show illustrations.) One of the brightest representatives of the frigate family is the frigate “Pallada”.
To be continued.
A fruit is that which arises from the gynoecium of a single flower due to fertilization or parthenocarpy. ( N.N.Kaden)
But not all types of fruits are formed from just one gynoecium. With the emergence of the inferior ovary and other forms of specialization of the flower, various parts of the latter take part in the formation of the fruit along with the gynoecium. Therefore, if the fruit is defined as a product of the development of the gynoecium, the concept of a “false” fruit must inevitably remain, a concept that is rejected by modern carpologists and, in particular, in the same work by Caden himself.
A.L. Takhtajyan was a proponent of a broad understanding of the fetus, interpreting fetus How the result of modification of the entire flower , and not just its gynoecium.
But even such a broad definition of the fruit does not include some of its types, such as sedge sacs, the fruits of some gonoceae, the fruit of ryegrass (a modified spikelet), etc.
The following definition seems more correct fetus :
A fruit is the gynoecium of one flower modified due to fertilization or apomixis, together with other organs of the flower and inflorescence that grow or remain with the gynoecium.
By adopting this definition of fruit, we will avoid the concept of “false” fruits. But at the same time, the sign of participation in the formation of the fetus must be introduced into the carpological classification adnexal organs, since the same type of fruit can be naked, with a veil, with a shell or wrapped.
According to Kaden - Takhtadzhyan fruit share based on the mutual fusion of carpels and the nature of placentation into the following main types:
I.
Apocarpies
- fruits produced from several free or one carpel.
II.
Cenocarps
- fruits produced from fused carpels. They include three types:
1.
Syncarpies- two- or multilocular fetuses with a central-angular placenta.
2.
Paracarps- single-locular fetuses with a wall placenta.
3.
Lysicarps- single-locular fetuses with a central placenta.
Caden divides each of these types into groups based on the number of carpels.
TO apocarpous fruits relate a lot of- And one-sheet; Various types boba, multi-nut and some drupes.
Syncarpies include syncarpous multileaflet; two- or multi-cavity boxes, fractional nut-shaped fruits “carceruli”(Borage and Lamiaceae); acorn And nut, and syncarpous drupes And berries.
Paracarps include paracarpous capsules(violets, willows, broomrapes); pods cruciferous, achenes Asteraceae, grains cereals and paracarpous drupes And berries.
Lysicarpous fruits relatively rare. Most typical boxes primroses and cloves; meet and lysicarpous drupes.
In addition to the basic carpological concept "fetus", it is necessary to clarify the content of such concepts as carpel, mericarp and fetal segment.
Under fruit you need to understand the part of the apocarpous fruit that is formed from one pistil.
Mericarp- this is a part of a syncarpous fruit (fractional), formed by one carpel or its longitudinal half.
Segment- this is part of an apocarpous (bean) or paracarpous (pod) fruit, disintegrating in a plane perpendicular to the longitudinal axis of the carpel.
In the literature devoted to the distribution of fruits and seeds, a distinction is sometimes made between morphological and biological understanding of the fetus, which do not always coincide.
Such a distinction between concepts should be discarded as methodologically erroneous. Any functioning organ has its own morphological nature, which mainly corresponds to the biological significance of the organ for the life of the organism. Therefore, the understanding of this or that organ in the morphological and biological sense is artificial. The erroneous concept of “fruit in the biological sense” arose due to the fact that when seeds are dispersed, not only seeds or fruits are separated from the mother plant, but also other parts of the plant that in one way or another contribute to seeding. Thus, the parts of plants with the help of which they (plants) reproduce and disperse are not always seeds or fruits.
The formations through which plant reproduction and dispersal are carried out are called rudiments. The term "primordium" corresponds to the terms "diaspora" or "disseminated", used mainly by foreign authors.
Rudiment may serve as a specialized cell, organ, part of an organ, or, conversely, a group of organs that are morphologically separate. In lower and higher seedless plants, the rudiment most often serves as a spore; in gymnosperms - a seed or sometimes a cone. The rudiments of angiosperms are the most diverse. According to their origin, the rudiments can be vegetative and generative.
Vegetative primordia of angiosperms- these are bulbs, tubers, sections of roots or rhizomes with renewal buds, etc.
Generative rudiments serve as seeds, fruits or infructescences.
Very often, the generative rudiment is a rather complex formation. For example, in some species of Lamiaceae, a calyx containing 1-2 or 4 “nuts” falls off. In linden (Tilia), as is known, the entire infructescence with the wing-shaped covering leaf of the inflorescence falls off. There are many similar examples that can be given.
The content also deserves special discussion. carpobiology(or otherwise - carpoecology
). In botanical literature it is commonly understood as carpobiology
ecology of distribution of primordia, including morphological adaptations to their distribution, in other words, The subject of carpobiology is considered just a process dissemination, i.e., dispersing seeds (fruits) in space. But this understanding of carpobiology is too narrow and leads to the fact that the process of dissemination is artificially separated from a number of phenomena closely related to it.
First of all propagation of primordia is happening not only in space, but also in time. The spread of primordia over time is due to the length of the dormant period of the seeds and their significant durability. All these phenomena are best studied in weed seeds. Research has revealed a remarkable relationship between the method of dispersal of primordia and the duration of their germination period.
Those rudiments that are adapted for long-distance dispersal (for example, equipped with flies) have a short dormant period and are characterized by amicable germination. Conversely, buds remaining close to the mother plant usually have a long dormant period. It is not uncommon for such morphological and physiological differences to be observed in the rudiments of the same individual.
These phenomena become understandable if we consider that the distribution of rudiments in time and space is equally increase the competitive capabilities of the species.
Secondly, it can often be observed that in the process of dissemination, the rudiments are prepared for germination. Thus, seeds or seeds of juicy fruits germinate better after they pass through the digestive canal of a bird or other animal. Some seeds weeds, collected from under a thresher, germinate much more vigorously than seeds of the same species collected from plants in the field.
In Mangroves, as is known, seed germination even precedes the separation of the primordia from the mother plants.
All these facts lead to the conclusion that the physiology of seed germination is closely related to the process of dissemination and is sometimes one of its aspects.
Third, the intensity and nature of the dispersion of the primordia are related to the period of their maturation and the duration of seeding. Thus, the ripening of juicy fruits at the end of summer and autumn is of great importance due to the fact that it is during this period that the number of insects decreases and birds switch to plant foods. Myrmecochory plants, on the contrary, bear fruit in late spring or early summer, that is, just when ants are especially active in collecting seeds. The number of such examples can be significantly increased.
Thus, carpoecology
should include the ecology of fruit and seed ripening, distribution, dormancy, and germination conditions. In short, subject of study carpobiology
there must be a biology of seed propagation as a whole.
Concerning ecology of propagation of primordia, then it is included in carpobiology as one of its sections.
Concept "device" is one of the most commonly used concepts in all biological literature, and most often under adaptation refers to one or another purposeful property (sign) of an organism. However, the question of the appropriateness of various devices must be decided specifically for each specific case. Darwin not only illustrated the relativity of the expediency of organisms with many striking examples, but also explained the inevitability of this fact: “Even if organic beings were fully adapted to environmental conditions at one time, they cannot remain so after these conditions change , unless they themselves change accordingly." Darwin draws our attention to another, no less significant side of the issue. He points out that in organisms, undoubtedly perfectly adapted to their place in nature, many aspects of organization do not have a direct and immediate relationship to modern living conditions. Let us add that we cannot judge the appropriateness of many adaptations simply because we do not know their biological significance.
How should we evaluate adaptations to dissemination: Are they always appropriate?
The overwhelming majority of works provide too little data on the actual dispersal of rudiments with the help of certain devices. Most often, only a priori, speculative conclusions are given about the meaning of these devices by analogy with other similar cases.
Under adaptations to the distribution of fruits and seeds
We will understand the morphological and physiological characteristics of the primordia and the entire plant that correspond to this method of dissemination.
Along with the concept of devices, it is necessary to touch upon another particular issue. In works on general carpology one can often find the expression: “fruits lacking adaptations for dissemination”. Although this idea is legitimate in morphological carpology, it should be taken into account that with a broad understanding of the fruit, there will be very few fruits that lack adaptations for dissemination; The various parts of the flower and inflorescence remaining with the gynoecium most often serve to distribute fruits or seeds.
At the same time in carpobiology
One cannot limit oneself to the study of fruits alone, since adaptations to dissemination are never limited to fruits and seeds alone. These adaptations cover the entire plant as a whole, its various organs and functions. Thus, by the time of seeding in herbaceous plants, the stem often lengthens, becomes elastic and rises upward. In other cases, it’s the other way around: the weak stem bends or even falls to the ground. The pedicels become elastic and bend in different ways, the calyx changes, etc. Of great importance for dissemination is fruit ripening period and seeding duration. Thus, not only the rudiments, but also the entire plant as a whole are adapted to one or another method of dissemination.
Basic special terms denoting various devices and methods of dispersing rudiments, often associated with the Greek root "choreo"- "spreading".
Plants in which the distribution of primordia occurs without the participation of any intermediaries or spreading agents are called autochoric, and the phenomenon itself - autochory
.
In contrast to autochoranes, they distinguish allochoric plants (“allos” - other), in which the rudiments are spread by some agents. Such spreading agents can be wind, water, animals and humans. The methods of dissemination corresponding to these agents are called anemochory
,
hydrochoria
, zoochory
And anthropochory
.
The distribution of rudiments by animals or humans can be carried out in different ways, therefore smaller categories are distinguished within the categories of zoochory and anthropochory.
In many plants, the dispersal of primordia occurs with the help of throwing devices. Such devices are called ballistic
(“ballo” - I throw, to the sword), and the plants that have them - ballistas
.
In the carpobiological literature one can often find the statement that the “simplest” method of dissemination is the abscission of the primordia under the influence of gravity ( barochory
). It is difficult to say what the authors mean by the words “the simplest”: the simplicity of the morphological structure or the phylogenetic primitiveness of barochory?
The idea of the “simplicity” of barochory is apparently based on two erroneous assumptions. Firstly, on the fact that there are supposedly fruits “devoid of adaptations for dissemination,” which is where barochors are included; and, secondly, on the generally accepted opinion that fruits with some appendages are always more perfect than fruits devoid of these appendages.
Those forms of barochory that are known in modern angiosperms should be regarded collectively as a perfect category of autochory, and in any case they are not “the simplest” in the phylogenetic sense.
Barochoria is most often characteristic of plants with single-seeded, indehiscent fruits, which, of course, are secondary to dehiscent multi-seeded fruits. Therefore, barochoras include many species of such families as goosefoot, buckwheat, madder, and grasses. Among legumes, for example, barochory is also observed in more secondary genera: sweet clover, some types of clovers and sainfoin. Barochory is very often (if not always) associated with heterocarpy, therefore, with the anatomical and physiological specialization of the primordia. Many barochors are inhabitants of specific plant groups: mangrove forests, deserts, crops. In each of these habitats, barochory is combined with peculiar features of dissemination: germination of unfallen fruits in mangroves; mucilage of seeds and synaptospermia in desert barochores; high fertility, long seed viability and extended period of germination in weeds.
Of all the forms autochoir the most primitive is, one must think, automechanochory
. The complex anatomical structure of the pericarp, which ensures the opening of the fruit and the scattering of seeds, is not biologically justified by the insignificant effect in terms of the range of seed dispersal. Automechanochory It is relatively rare and is characteristic mainly of families that occupy a low position in the system: moth, geranium, oxalis, balsamaceae. In more secondary families - violets, euphorbias - mechanochory is combined with myrmecochory, which plays a major role in seed dispersal here.
The most specialized form autochoir certainly is geocarpy
, in which there is a sharp deviation from the typical fruiting pattern for all plants. The formation of underground fruits, characteristic of geocarpic species and forms, is, of course, a secondary phenomenon that arose as an adaptation to the peculiar conditions of deserts and semi-deserts. Basicarpic and amphicarpic species represent quite obvious links between plants with terrestrial fruits and geocarpic species.
Geocarpy
- the most specialized method of dissemination not only among autochorants, but generally among all natural methods of dissemination of rudiments.
Among the varied and numerous ballistic devices We distinguish four types: Carnationaceae, Lamiaceae, Umbellaceae and Asteraceae.
The largest and undoubtedly the most primitive of all these types is the type of carnation, represented by a variety of opening multi-seeded fruits. In a genetic sense, the type of cloves, of course, is a composite one, therefore, within the type, it is necessary to distinguish ancient representatives from relatively young and progressive ones.
The most primitive type of ballistic fruit, not only in the “carnation type”, but in the entire category of ballistae, is dry multi-seeded multileaf
, for example, in Trollius europaeus. Very close to her three-leaflet
Aconitum and Delphinium. Ballistas must be recognized as secondary in this type boxes bellflowers and orchids with their peculiar method of opening, formed from the lower ovary.
The ballistae of the “Lamiaceae type” are more advanced compared to the previous type, if only because the rudiments here are not seeds, but indehiscent single-seeded mericarps and less often - fruitlets; in addition, it takes part in the dispersion of rudiments cup. The simplest representatives of this type of ballistae are cinquefoils, whose fruit is multi-nut , and the calyx does not undergo any specialization. Secondary species should be considered those species of Lamiaceae that have a zygomorphic calyx that changes significantly after flowering and during fruiting (Prunella, Scutellaria and others).
In terms of the nature of their ballistic devices, the phylum Compositae is close to the Lamiaceae. Higher specialization ballistas of the Asteraceae is manifested in the fact that they the entire basket functions as one multi-seeded fruit.
The most unique of all ballistic devices is carpophorus umbrella Vislopordnik the umbrella type is extremely specialized and represents a secondary formation, as it originates from the lower syncarpous drupe. Thus, the most primitive ballistic adaptations among angiosperms are represented in the multileaf type of swimsuit, and the most secondary ones are in the umbelliferous umbelliferous and the Asteraceae basket, which functions as a single multiseeded fruit.
Ballistic devices, like any other adaptations to dissemination, arose many times at various stages of angiosperm evolution. In other words, different types of ballistas are characterized by phylogenetic diversity in age, but in most cases they can be qualified as a progressive carpoecological category. The following features inherent in ballistas lead to this conclusion.
Firstly, among the seed plants, ballistas are only herbs, therefore angiosperms.
Secondly, the category of ballistae is widespread in nature, especially in steppe phytocenoses, which are secondary in relation to the tree type of vegetation.
Thirdly, a common feature of all ballistae is the absence or minimum of neoplasms associated with this method of dissemination.
Ballistic adaptations come down mainly to the lignification of stems and peduncles, which is generally characteristic of most fruit-bearing plants; to a change in the position of pedicels or sepals during fruits, etc. In short, ballista devices are distinguished by their simplicity and “economical” structure. This is especially clear when comparing ballistae with many anemochores, epi- or endozoochores with their bulky flying or tenacious appendages, with massive succulent tissues.
The simplicity of the structure in many cases is undoubtedly primary and indicates the primitiveness of this structure, but when we are faced with fruits of a clearly secondary type, as in Lamiaceae, Umbelliferae, Compositae, etc., the simplicity of the adaptations should already be considered as a result of simplification. In these cases, Timiryazev’s position is true: “The solution to the same problem in the simplest ways should be recognized as a sign of perfection”. This requirement is precisely what ballistas satisfy: the problem of dissemination is solved by them in the “simplest ways” in the sense of simplicity of devices.
Finally, ballistas are widely represented in such young families as borage, labiaceae, noricaceae, primroses, umbellaceae, bellflowers, and asteraceae. Anemochory with dusty seeds in the families of wintergreens, broomrapes, orchids and others are, at the same time, clearly expressed ballistas. Likewise, in families where ballistas are found along with other categories, the former belong to the more progressive genera.
In the family of moths, ballistae are known among astragalus and octopus, the fruits of which are certainly secondary than ordinary beans characteristic of auto mechanic choirs
.
Among the umbrellas in the group Hydrocotyloideae, which is unanimously considered more primitive by all taxonomists, there is no free carpophora. Thus, ballistae in the Umbellaceae family are also derivative, more advanced forms.
It is easy to prove that in the Asteraceae family, ballistae are secondary to anemochores.
There are a number of facts indicating a reduction of the crest in some evolutionary series of Compositae. However, reduction of the tuft in this family most often leads to ballistic dispersion of the fruits. In addition, in Ballistae asteraceae, the entire basket as a whole is adapted to dissemination and represents, as it were, a single fruit, whereas in anemochores the pappus ensures the flight of only one achene.
If the inflorescence of the Asteraceae, functioning as a single flower, serves as the most important argument in favor of the progressiveness of the family, then with the same reason the infructescence functioning as a single fruit is an indicator of high specialization.
Finally, the biological perfection of ballistae is manifested in the fact that for them, any factor capable of shaking the stem can serve as an agent of dissemination: wind, animals, agricultural implements, various types of transport, or sometimes pedestrians.
Zoochoria
in seed plants it is represented by a greater variety of adaptations than anemochory. At the same time, the phylogenetic diversity of ages of various forms of zoochory appears even more clearly.
The most ancient type of seed plant rudiments are seeds with a succulent seed coat. This is evidenced by the rudiments of cycads and ginkgos - the most ancient representatives of gymnosperms. As studies have shown, anatomical structure The magnolia seed coat also bears the imprint of great antiquity. Type of magnolia seed coat Russian botanist K.K. Zazhurilo is considered as a source not only for the family Magnoliaceae, but also for all angiosperms. The same author established great similarities in the anatomy of magnolia seed coats with cycads and ginkgos. Regardless of whether this similarity is homologous or only analogous, it gives reason to assume the same method of seed dispersal in the named plants.
Thus, endozoochory should be regarded not only as the most ancient form of zoochory, but also as the most ancient method of dissemination of seed plants in general.
A comparison of data from paleobotany and paleozoology suggests that the first agents of seed dispersal of ancient gymnosperms and magnolias were arboreal forms of reptiles, in other words, at the dawn of the evolution of angiosperms, only endosaurochory.
Another ancient type of endozoochorous primordia, characteristic only of angiosperms, is juicy flyer. It is present in Schisandra, most Anonaceae, and is occasionally found in Ranunculaceae.
But it would be a grave mistake to say that endozoochory in general and, therefore, any endozoochorous rudiments are primitive. Genetically different types of juicy fruits, from apocarpous drupes to paracarpous berries, arose many times during the evolution of angiosperms in a variety of directions and at different stages of their development. This is why juicy fruits, not to mention their morphological nature, are found in the families Berberidaceae, Rosaceae, Caprifoliaceae, Rhamnaceae, Rutaceae, Vitaceae, Ribesiaceae, Solanaceae, Vacciniaceae, Araliaceae, Liliaceae, Araceae and many, many others.
In modern genetic classifications of fruits of various types, juicy fruits are considered as derivatives of dry ones: the apocarpous drupe comes from the leaflet; syncarpous and paracarpous drupes and berries - from the corresponding types of boxes. Anatomical and carpological studies by K.K. Zazhurilo established in the pericarp of juicy fruits rudimentary structures of opening mechanisms of capsules, which are the initial type of fruit for the studied berries and drupes. According to K.K. Zazhurilo, traces of origin from capsules are found in the juicy fruits of Daphne altaica, Cucubalus baccifer, Asparagus officinalis, Phellodendron amurense, Hypericum sp. and etc.
The emergence of juicy fruits from dry ones in the various evolutionary series of angiosperms undoubtedly went in parallel with the vigorous formation of birds, which took place in the Tertiary period. In the Cretaceous, most forms of birds apparently belonged to aquatic flightless birds. At the beginning of the Tertiary period, flying forms occupy a central place. In the Middle and Upper Eocene, most modern orders of birds were already represented; passerines are observed for the first time. Ho modern species There were no birds yet in the Miocene. They appear in the Pliocene, and finally, in the Pleistocene, the composition of the bird fauna basically becomes quite modern.
One must think that further evolution endornitochory manifests itself in the confinement of succulent species to certain phytocenoses, and on the other hand, in the selectivity to food of frugivorous birds. As a result of this specialization, types of juicy fruits will be selected that are capable of fully maintaining seed germination in the digestive canal of certain bird species. How can one imagine such an extreme degree of specialization? adaptability of certain types of fruits to bird species, as is observed in some flowers in relation to insect pollinators.
In certain evolutionary branches there was transfer from endozoochory to other methods of dissemination. In the buckthorn family, for example, one can trace the transition from the upper succulent drupes to the dry winged or fractional drupes developing from the semi-inferior and lower ovary.
Endozoochorous distribution of dry buds plays a much smaller role in nature than the distribution of juicy fruits and seeds. Some features of the endozoochorous distribution of dry primordia allow us to conclude that this form has a secondary nature zoochoria
.
First of all, the main agents of endozoochorous spread of dry fruits are herbivores mammals, in particular livestock
. In this regard, this method of dissemination is characteristic of herbaceous plants of steppes and pastures and weed-ruderal species.
Secondly, the successful endozoochorous distribution of dry primordia is associated with their physiological resistance against digestion, which is already a sign of specialization. Finally, very often endozoochorous species with dry primordia are characterized heterocarpy, thanks to which this method of dissemination turns out to be the most effective. The secondary nature of heterocarpy has already been discussed above.
What evolutionary assessment can be given? epizoochory?
In a relationship facultative epizoochory , i.e. spreading of germs with soil and silt on the legs and body of animals, it is difficult to make any phylogenetic assumptions. This method of distribution of rudiments is determined not so much by the properties of the rudiments themselves, but by the habitat conditions of the species.
Concerning epizoochoric spread of tenacious fruits (specialized epizoochory
), then we can assert the secondary nature of this method of dissemination: tenacious fruits or seeds are known only in herbs and are completely absent among trees and shrubs. Tenacious seeds are very rare. Typically, tenacious and spiny appendages develop on single-seeded, indehiscent fruits, therefore epizoochory is known in such undoubtedly progressive families as borage, madder, umbelliferae, and Asteraceae.
Spiky and tenacious fruits develop mainly in plants of two environmental groups: in the inhabitants of dry and hot steppes and deserts and in weed-ruderal plants. These habitats are secondary.
The most progressive form zoochorous propagation of primordia should be considered synzoochory
and in particular, myrmecochory
.
Sinzoochory
is associated with a complex instinct of agents of dissemination - with the instinct of storing food and with a rather strict selectivity for food.
Synzoochorous rudiments, distributed by rodents and birds, are clearly of a secondary nature. These are either specialized coniferous seeds such as " pine nuts", or dry single-seeded fruits of angiosperms: various types of nuts and acorns, which Takhtajian defines as "very specialized types of lower syncarpous fruit."
Secondary features appear even more clearly myrmecochory
. If synzoochorous rudiments distributed by rodents and birds serve as the basis of food for them and a significant part of these rudiments is destroyed, then myrmecochorous fruits and seeds are not eaten by ants. Myrmecochorous primordia are protected by strong integuments and, as is known, are equipped eliosome- a special appendage that serves as bait and food for ants. Besides, myrmecochory
characteristic only of herbaceous plants and most often of progressive families, such as Scrophulariaceae, Violaceae, Aristolochiaceae, Euphorbiaceae, Labiatae, Boraginaceae, Santalaceae.
Myrmecochory
extremely widespread in nature in all geographical latitudes and plant groups, with the exception of the far north and swamps. Finally, it should be added that ants are among the most secondary groups of insects.
These are the main paths of evolution zoochorous methods of dissemination from endosaurochory to myrmecochory.
Among the various anemochorous adaptations the most primitive are pterygoid appendages on fruits and seeds. Winged seeds are known from many conifers, including such ancient ones as araucaria and sequoias. Winged fruits are characteristic of anemochorous magnolias (tulip tree) and some other relatively primitive families (Dipterocarpaceae). And among other families of angiosperms, winged primordia are most often characteristic of trees and rarely of grasses.
Of all types of wing-shaped formations the most perfect seems single-sided asymmetrical wing with heavily reinforced leading edge, i.e. the wing of a maple fly. Wings of this type impart screw flight to the rudiments, as a result of which, falling from a height even in a very weak wind, the fruits are carried away to a relatively long distance.
Hair-like appendages found only on the fruits and seeds of angiosperms and almost exclusively in grasses. From this we can conclude that the hair fly is phylogenetically younger than the wing, although it is not its derivative.
The simplest form parachute present hairs covering evenly the entire rudiment(anemone, cotton grass, etc.). Such a fly is not very effective, since the buds are usually linked by their hairs and form loose clumps that weakly resist the air flow.
Hairs at the base of the primordia or on their sides also did not justify their worth in the course of evolution: such parachutes are very rare.
The most common type of parachute is tuft of hairs at the apex of the bud. This one fact alone suggests that this type of parachute is more advanced than others. The pappus at the top of the fruit or seed ensures stable balance of the rudiment in the air, since the center of gravity of the rudiment is below the parachute. Such rudiments are well adapted for soaring flight. They are characteristic of secondary families, such as the Palmaceae, Valerianaceae, and Asteraceae.
The higher the parachute is raised above the center of gravity of the rudiment, the more stable the balance such a rudiment maintains in the air. That's why Fruits in which the crest is attached to an elongated nose are best adapted for soaring flight., especially if the crest consists of feathery hairs and has stronger resistance to air flow. All these characteristics are possessed, for example, by the achene of the greater salsify, in which the spout is equal in length to the achene and is crowned with a feathery pappus reaching 4-5 cm in diameter.
Thus, the entire evolutionary path of feathery anemochoric primordia can be outlined from the nut of the anemone to the achene of the greater salsify.
The other two types anemochoric primordia - small dusty seeds
And “balloon” fruits
- seem to be even more secondary than the rudiments with a feathery crest.
These facts lead to the conclusion that anemochory , associated with a very low weight of the primordium, is a secondary phenomenon for angiosperms.
“Balloon” fruits
, well adapted to anemogeochory, are found in various families. The nature of the adaptations in such fruits is also different (swollen cups or bean valves, meridionally located wings, branched elastic appendages, etc.), but they all have one common feature: streamlined shape combined with low specific gravity. This type of anemochora is confined mainly to sandy deserts and is an ecologically specialized group of plants.
Analyzing the formation of the vegetation cover of the hot deserts of Central Asia, it is noted that the genera Colligonum and Smirnovia are progressive, and the swollen sedge (Carex physodes) is characterized as an example of “shape formation of the current day.” All these plants have fruits of the type "balloon".
The study of the anatomy and ecology of some types of swollen fruits has shown that the structure of their pericarp is essential for seed germination, and perhaps anemochory is only a consequence of this structure. Thus, the fruits of different species of Prangos and Cachrys from Umbelliferae have very light fruits with a relatively large volume (100 mg with a volume of 1.4 cm3). The pericarp tissue of these fruits is very similar to the velamen tissue of the aerial roots of epiphytic orchids. From this we can assume that the pericarp serves to absorb and retain moisture from the air. Indeed, experiments have shown that Prangos fruits absorb 33% of their dry weight of vaporous water in 7 days, and Cachrys fruits - 20%. It is remarkable that the fruits of different Prangos species absorb more water, the more xerophytic their habitat.
In passing, we note that the winged fruit of the tree (Paliurus) ,
thanks to its thin cuticle and loose mesocarp, “it has the ability to absorb moisture over its entire surface, like a sponge.”
Generally anemochory is regarded by many botanists as a progressive character, indicating a higher position in the system of anemochoric genera and species, in comparison with non-anemochoric ones. It is impossible to agree with such an assessment of anemochory in general, regardless of the nature of the adaptations.
First of all, we saw that various anemochorous adaptations arose at different stages of the evolution of seed plants, therefore some of these adaptations - wing-shaped appendages - are already known in ancient conifers and primitive angiosperms.
In addition, in a number of cases, anemochorous rudiments are certainly not the most progressive within the family or genus. Thus, studying the anatomy of the seed coats of the Magnolia family, Zazhurilo comes to the conclusion that among the Magnoliaceae, three types of seed coats can be distinguished, with the type Liriodendron (anemochore) occupying an intermediate position between the more ancient type Magnolia and the most progressive Schisandra (endozoochory and other methods).
Fruits of Pastinaca sativa and Heracleum sp. of the umbelliferous ones have a clear anemochoric character (strongly flattened with a wing-shaped border), but the ontogeny of these fruits and the nature of the mechanical layer of the pericarp indicate their closeness to Hydrocotyloideae- the most primitive group among the umbelliferae.
We have already dwelled in detail on the phenomena of crest reduction in the Asteraceae family. It can be argued that many non-anemochorous Asteraceae are certainly secondary to anemochorous ones.
Finally, in some cases, fetal characteristics that are anemochorous in nature may have a completely different adaptive significance, as is observed in various types Prangos and Paliurus .
Thus, anemochory at all cannot be regarded as an unconditionally progressive sign. Various forms of anemochory in different families and genera are either primitive or secondary. In some evolutionary series, there is a loss of anemochorous adaptations and a transition to other methods of dissemination.
General characteristics of the fetus. The fruit is a characteristic organ of angiosperms. It is formed as a result of the changes that occur in the flower after fertilization. Sometimes the fruit is defined as a mature flower. The most essential part of the fruit is the gynoecium, however, in many plants, mainly those with an inferior ovary, other parts of the flower, primarily the receptacle and peduncle, also take part in the formation of fruits , and sometimes parts of the inflorescence. The fruit largely retains the characteristics of those parts of the flower from which it arises, but the original structures often undergo profound changes. Therefore, in the structure of the fruit, along with the signs of the gynoecium and other parts of the flower, there are signs of the fruit itself, which distinguish the fruit from the corresponding parts of the flower very significantly. Only in the simplest cases (buttercups, legumes) do mature fruits differ from the gynoecium only in size; often they take on such unique features that it is difficult to determine from which gynoecium they arose.
The diversity of fruits is determined by three groups of characteristics:
the structure of the pericarp;
method of opening or disintegration;
features associated with distribution.
Pericarp. The pericarp is an overgrown and often highly modified wall of the ovary, which, together with other organs of the flower, is included in the composition of the fruit. In the pericarp there is an outer layer - the exocarp and an inner layer - the endocarp, and sometimes a middle layer - the mesocarp. It is not always easy to distinguish these zones. All three zones can be most clearly distinguished in drupe type fruits - a thin leathery exocarp, a fleshy mesocarp and a hard endocarp. In typical berries, the entire pericarp is juicy and individual layers are difficult to distinguish. Also, in dry fruits, in some cases one can detect layers of differently differentiated cells (sunflower), while in others the pericarp is completely homogeneous (hazel).
During the process of ripening, the pericarp undergoes very significant biochemical changes, the accumulation of sugars, vitamins, various aromatic substances, fats, etc. occurs, on which the use of fruits by humans and animals is based. The pericarp of mature fruits, as a rule, no longer contains chlorophyll-bearing layers. The fruits become brown or acquire a bright color due to the formation of carotenoids, anthocyanins, etc. Not only juicy fruits, but also dry fruits, for example, some maples, are brightly colored.
Principles of fruit classification. The defining morphological feature of the fetus is the type of gynoecium from which it develops . In connection with sapocarpous, syncarpous, paracarpous and lysicarpous types of gynoecium, fruits are distinguished as apocarpous, syncarpous, paracarpous and lysicarpous. Each of these types is divided into subordinate groups, also in connection with the main directions of the evolution of the gynoecium. Among apocarpies, a distinction is made between polymeric, i.e., arising from several or many carpels, multi-seeded and single-seeded and monomeric multi-seeded and single-seeded fruits. In each of the coenocarp types, one can distinguish between upper and lower multi-seeded and single-seeded variants. Finally, in each of the latter there may be fruits that differ in the method of opening and spreading.
In some cases, artificial classification of fruits is also possible, based mainly on signs of external morphology. All fruits are divided into juicy and dry. The latter are divided into dehiscent and non-dehiscent, etc. Moreover, one term often denotes completely different formations. Thus, nutlets are the fruits of buttercups arising from one carpel and the fruits of birch, formed from the coenocarpous gynoecium at the lower ovary. How boxes denote various types of dry dehiscent fruits that do not fit other terms, etc.
Opening the fruits. Opening refers to the release of seeds before they germinate. In this case, special separating tissues are formed in certain areas of the pericarp (pericarp). In the vast majority of cases, it is clear that dehiscence is characteristic of dry polyspermous fruits. In other cases, the pericarp is gradually destroyed by mechanical influences, the activity of microorganisms, etc. Such fruits are called non-dehiscent. Most often, opening occurs through longitudinal slits. In this case, the cracks appear along the ventral suture (ventricidal dehiscence), along the midrib of the carpel (dorsicidal dehiscence) or along the surface of the carpel (laminal dehiscence). In syncarpous fruits, rupture along the septa may occur, i.e. in the plane of carpel fusion. In this case, they speak of septicidal fruits, in contrast to loculicidal ones, which open in nests. In coenoparacarpous and coenolysicarpous fruits, opening can occur at the places where the carpels grow together (cruciferous, poppy, cloves) or along the midrib of the carpels (violets, willows). The longitudinal opening of the fruits can be complete - with valves - or incomplete, for example with cloves, which is especially typical for cloves. Incomplete opening can also include opening with holes (maki). Typically, incomplete dissection is a more progressive method than complete dissection, and can be derived from the latter. Sometimes a very peculiar cross-ring opening occurs (henbane, amaranth). It is usually accompanied by the formation of a cap .
Drawing – different opening of the boxes: 1 – cloves (primrose), cap (henbane), 3 – flaps (datura)
Decaying fruits are also, as a rule, more evolutionarily developed compared to dehiscent ones. They are divided into two groups.
Belonging to the first, the so-called fractional fruits, disintegrate longitudinally, in the plane of fusion of the carpels. In this case, closed single-seeded mericarps are formed, for example, two in number, as in the extensive umbelliferous family. In milkweeds, their fruits, formed by three carpels - “three nuts”, disintegrate in the same way, but in them the pericarp is torn and the mericarp is open on the ventral side.
A special type of disintegrating fruits from four nuts is characteristic of Lamiaceae and borage with a gynoecium formed by two carpels. The mentioned variants of the first group include coenocarpous fruits.
The second group includes the so-called articulated fruits, which include both apocarpous and coenocarpous variants. They disintegrate transversely in planes perpendicular to the longitudinal axis of the carpel. In this case, the segments usually remain closed due to the formation of transverse false partitions between them. Jointed fruits are especially characteristic of plants inhabiting areas with arid climates
Apocarpous fruits. Apocarpous fruits are represented mainly among polycarpous plants, as well as in such large groups as Rosaceae and legumes, in some primitive monocots and other plants. The primitive type of fruit, perhaps one of the original for angiosperms, is the multileaflet, which is a collection of leaflets.
A leaflet is a dry multi-seeded fruit formed by one carpel and dehiscent on one side. The number of carpels in a flower corresponds to the number of leaflets included in the multileaflet. Leaflets are opened ventricidal or dorsicidal.
Reducing the number of seeds in each fruit to one leads to the formation of a polynut, characteristic of many ranunculaceae, Rosaceae (for example, cinquefoil) and some monocots (chastuha). In the buttercup, which has a convex receptacle, the nuts cover approximately half of the surface of the latter. Placing a large number of nuts is possible thanks to the narrow bases.
It is important to note that while the leaflets remain on the receptacle, the nuts separate when ripe. Thus, the decrease in the number of seeds also resulted in changes in other respects. In some southern species of buttercups, the stylodes preserved in the fruits serve as attachments promoting zoochory (see below).
Figure – multi-nut gravilata: 1 – general view, part of the fruitlets has been removed, the receptacle is visible; 2 – carpel during flowering, 3 – nut, the lower part of the stylode is preserved; VSt is the upper part, NSt is the lower part of the stylodium; Tsvl – receptacle; G – gynophore
Strawberries should also be considered multi-nuts.
Despite the external difference, the fruit of the rose hip (Rosa), called cynarodium, is quite close in essence to the fruits of strawberries. The main difference is that instead of a convex axis, there is a goblet-shaped hypanthium of mixed origin (a floral tube formed by the fusion of the main lower parts of the perianth and staminate filaments; it looks like a concave receptacle. The main form is common in plants of the rose family ( rosehip, cherry, burnet, etc.), as well as in some tropical families), in the lower part of which nuts are attached; long columns protrude out through the narrow opening of the hypanthium.
The original type of polynut is found in the Indian lotus, which forms dense thickets, for example, in the Volga delta. The receptacle of the lotus grows greatly and each fruit is immersed in a special depression.
Drawing – Multi-nut of rosehip and Indian lotus
1 – longitudinal section through a rose hip flower; 2 – rose hip nut, 3 – Indian lotus fruit
Polydrupes and drupes are also often found among apocarpous single-seeded juicy fruits. Various representatives of the genus Rubus (raspberries, blackberries, etc.) have polydrupes. The fruit of these plants consists of many small drupes located on a common receptacle. Each drupe has a succulent, fleshy mesocarp and a stony endocarp, which contains a single seed. The number of drupes varies from 3-6 for drupes to several dozen for raspberries and blackberries. When ripe, raspberry polydrupes are easily separated from the convex receptacle, which looks like a white “stump,” while the fruits of blackberry species, especially numerous in the Mediterranean, where they often form impenetrable thickets, are separated along with the part of the receptacle to which the fruitlets are attached.
Monodrupes are well known, since they are possessed by all representatives of the plum subfamily of the Rosaceae family, which are very widespread in cultivation (the so-called stone fruits). In drupes, the differentiation of the layers of the pericarp is especially clearly manifested. The structure of the bones is especially varied, including the sculpture of their surface.
In cherries and cherries, differentiation into mesocarp and endocarp begins at the flowering stage, when both ovules are clearly visible, one of which then dies. Already in a young fruit the ovule fills the entire space inside the endocarp . IN At this time, the latter completely hardens, while the mesocarp is still far from fully developed. The abdominal suture is clearly visible in the form of a groove
Another direction of leaflet evolution led to the emergence of the bean. The bean differs from the leaflet mainly in that it opens both along the ventral suture and along the midrib of the carpel, i.e. dorsoventricular. In addition, the bean always arises from the monomeric gynoecium. The specialization compared to the leaflet is that when the bean ripens, the valves can instantly curl and scatter seeds, which is facilitated by special strands of mechanical fibers in the mesocarp. A dry polyspermous bean that opens with two valves is characteristic of most moths (astragalus, peas, lupine, gorse, etc.).
In some moths, as well as in the closely related families Mimosa and Caesalpiniaceae, the pods are specialized in connection with the mode of distribution. Thus, in the camel thorn, which lives in desert areas, the beans disintegrate into segments. The spirally twisted and often still thorned fruits of some alfalfas are unusually unique. , living in dry areas of the Mediterranean and Central Asia. In the same places there is also a Colutea shrub with bright yellow or orange flowers, large bubble-like swollen beans and very big amount ovules . The lateral branches of the ventral bundles are clearly visible. It is also worth noting the uniqueness of clovers, in which the beans are surrounded by perianth even in a mature state; opening is obviously a rudimentary function for them. The tropical vine Entada, a member of the Mimosa family, has the longest fruits in the plant kingdom. Its beans are 10-15 cm wide and reach one and a half meters in length.
Figure - Variety of beans and the carpel of legumes: 1 - camel thorn, 2 - alfalfa, 3 - bladderwort, 4 - bladderwort carpel after flowering, opened on the dorsal side
A reduction in the number of ovules leads to the appearance of single-seeded beans that do not open like nuts (many clovers, some gorse, alfalfa, sainfoin).
Syncarpous fruits. The most primitive syncarpous fruits - syncarpous multileaflets - are characterized by opening in the area of the upper free sections of the carpels. They are still very close to apocarpous fruits. Such fruits are characteristic, for example, of black buttercup (Nigella) from the Ranunculaceae. With complete fusion of carpels, the number of which is often impossible to determine by appearance fruit, a fruit appears - a syncarpous capsule with several nests corresponding to the number of carpels involved in the formation of the gynoecium. Capsules are present in species of the most diverse families of dicotyledons and monocotyledons, but arise predominantly from the superior ovary. The methods for opening boxes are quite varied. A good example of a syncarpous capsule is the fruit of a tulip.
Figure – Syncarpous tulip box: 1 – general view before opening; 2 – transverse section through the ovary; SP – vascular bundles
Among the syncarpous fruits, the so-called fractional fruits are extremely diverse, breaking up along partitions into mericarps (the single-seeded portion of the fruit), corresponding to individual carpels. In the simplest cases, they are still close to opening fruits (in milkweeds). Also well known are fruits that fall apart into open single-seeded lobes - malvaceae cakes. In this case, opening, as such, often does not occur.
The maple tree, equipped with two wings directed to the sides, also belongs to the fractional fruits. .
Fractional fruits are formed from the lower and semi-inferior ovaries. Thus, in bedstraws and woodruffs from the madder family, the fruits split into 2 more or less spherical mericarps, smooth or with attached
Peculiar and characteristic fruits, the so-called elliptical fruits, are formed by representatives of the large family of Umbelliferae
Drawing – fractional fruits: 1 – maple diptera; 2 – bedstraw fruit; 3 – the same after separation of the mericarp; 4 – umbelliferum
Finally, a special type of decaying fruit is represented by coenobia, characteristic of the species-rich borage and labiatae.
In coenobia arising from the superior ovary, decomposition occurs not only along the septum between two carpels, but also along an additional septum perpendicular to the latter. The distribution unit thus corresponds not to the carpel, as in the variants described above, but to its half; from the ovary, consisting of two carpels, 4 “nuts” are formed. The reflexing lobes of the calyx and various attachments also contribute to the distribution of fruits of many borage species.
There are many plants that form syncarpous juicy fruits. The upper syncarpous berry is known, for example, from grapes, crow's eye, potatoes, tomatoes, etc. Here, unlike drupes, the entire pericarp is juicy, and the seed coat is hard, containing stony cells.
Special mention should be made of the tomato fruit. Like other nightshades, the ovary here is bilocular, formed by two carpels with a large number of ovules. But cultivated varieties, as is well known, are characterized by a multi-chambered fruit, which arises as a result of an increase in the number of carpels due to the fusion of several flowers. Not only the pericarp, but also, to a large extent, the placentas, which grow to such an extent, participate in the formation of the juicy pulp of the fruit - the pulp that the seeds are immersed in their mass. The seed coat, which is subject to mucus, also undergoes significant changes.
The upper juicy carp of citrus fruits, the so-called hesperidia, are very peculiar. The orange ovary is multilocular, coenosyncarpous, with central-angular placentation. The thickness of the ovary wall with relatively thin septa is noteworthy. The pericarp developing from the wall of the ovary is differentiated into a dense, leathery exocarp with a large number of essential oil, painted in yellow carotenoids, n spongy white mesocarp - albedo. The edible pulp of the fruit - the pulp - is a remarkable new formation resulting from the fact that outgrowths begin to appear on the inside of the pericarp, developing into juicy sacs, gradually filling the ovary nests and growing between the seeds . The albedo is a degenerating tissue and therefore in the mature fruit the pericarp is easily separated from the pulp.
Figure - Orange fruit: 1 - diagram of a cross section through the ovary; 2 – fetal sector in cross section; A – albedo; C – seeds
Juicy syncarpous fruits arising from the lower ovaries are common in species of the lingonberry and honeysuckle families. Lingonberries, blueberries, and honeysuckles (sometimes called wolfberries) produce berries, and elderberries produce multi-stone drupes. The well-known tropical fruits of the coffee tree and bananas are also located here. The coffee tree belongs to the madder family, just like our bedstraws. All members of the family are characterized by an inferior ovary. However, unlike our madder fruits with dry fruits, coffee fruits are drupes. Banana fruits belong to the lower syncarpous berries, although they bear little resemblance to berries in the usual sense . Their exocarp is leathery and relatively thick, the inner layers form a juicy powdery pulp. Bananas, known to everyone, are cultivated seedless forms. As can be seen from the diagrams , The ovary of a wild banana is very similar to the superior ovary of a tulip.
A special place is occupied by the apple fruit, which represents an example of specialization at a rather low evolutionary level. Fruits of this type are characteristic of the apple subfamily of the Rosaceae family, in particular apple, pear and quince. In a cross-section, 5 single-seeded nests are visible through the apple.
The parchment-like wall of each nest corresponds to a carpel, of which, therefore, there are also 5. Since the carpels do not grow together with each other, some scientists classify the apple as an apocarpous fruit, but others believe that the parchment-like part is only an endocarp, and the outer tissues of the carpel become fleshy and completely merge with the tissues of the flower tube.
Figure - Apple tree: 1 - cross section, 2 - apple with carpels removed, Pl - border of carpels
Finally, the fruits of the pomegranate, the only species of the family, are very specific. The pomegranate fruit develops from the lower ovary and has a dry, leathery pericarp that opens in irregular cracks when ripe. The nests are filled with large seeds with a bright red, garnet-colored, very juicy peel; the latter, as already mentioned, represents a very rare case.
The syncarpous fruits considered are multi-seeded formations. Next we will talk about single-seeded syncarpous fruits. They are also quite numerous and develop from both the upper and lower ovaries. The first includes, for example, the “coconut”, reaching a length of 30 cm and a width of 20 cm. The seed - one of the largest in general - initially has a liquid endosperm, which is consumed as the so-called “coconut milk”.
Among the lower one-seeded syncarps, the most famous are the fruits of various catkins (often called nuts, which, of course, is incorrect, since the nut is an apocarpous fruit). A very specialized fruit is the nut - the fruit of the hazel, or hazel. Since the ovary here has two stigmas, it can be assumed that it is formed by the same number of carpels. It is bilocular, syncarpous, with one ovule in the nest ; During fetal development, however, the septum is transformed into a column and only one ovule develops. The young nut has a powerful pericarp , Moreover, the exocarp is sclerified, and the mesocarp is a spongy tissue that fills almost the entire internal space. Subsequently, it degenerates and the vacant space is filled with a single developing seed.
Figure - Hazelnut: 1 - diagram of the ovary, 2 - the same at a later stage, 3 - longitudinal section through the unripe fruit; S - seed, K - column, ASz - removed ovule, End - endocarp
The oak acorn differs from the nut in having a leathery, non-woody pericarp and a different origin of the acorn. The latter, in the early stages of development, almost completely surrounds the ovary. Subsequently, the plush lags in growth from the fruit, which in its mature state protrudes significantly above the edge of the plush . Three stigmas indicate that the fruit is formed by three carpels. Each carpel contains 2 ovules, but, except for one, all the others are reduced. After removing the pericarp, 6 reduced ovules can be seen in the lower part of the mature fruit .
Paracarpous fruits. Similar to what was stated above regarding the gynoecium, paracarpous fruits could have arisen from syncarpous ones or formed in the early stages of the formation of angiosperms from unfused carpels. Paracarpous fruits are characteristic of species of different families of angiosperms, so they undoubtedly arose in different evolutionary lines; among the paracarps there are polyspermous and single-seeded, dehiscent and indehiscent, upper and lower.
The initial types of paracarpous fruits include paracarpous boxes and pods. The former are common, for example, in species of the poppy family (celandine fruit).
Figure – Paracarpous capsule of celandine: 1 – unripe fruit, 2 – dehiscent fruit, P – frame
A special type of lower paracarpous fruit is formed in species of the pumpkin family. The pumpkin fruit is characterized by a hard, often very strong exocarp and a fleshy mesocarp. The fetal cavity is filled with placentas, often very juicy. In some pumpkins, high osmotic pressure arises in the cells of the inner layers of the pericarp during ripening, causing the fruits to open and the seeds to be scattered over a fairly considerable distance (several meters in the crazy cucumber - Ecballium elaterium).
In addition to some cruciferous plants, single-seeded upper paracarps are characteristic of species of a number of other families, primarily such large ones as sedges and grasses.
The grain of cereals is an indehiscent single-seeded fruit, in which the thin pericarp is so closely adjacent to the seed coat that it seems fused with it. In most types of cereals, the caryopsis falls off along with the scales surrounding it. Naked grains are found only occasionally in wild cereals, but are quite common in cultivated varieties. Various types of appendages are often formed on the flower scales, facilitating the spread of fruits. The feathery awns of feather grass are especially hygroscopic, being several times longer than the grain itself. Some cereals from the bamboos group produce berry-shaped caryopsis.
The lower paracarpous fruits include the fruits of Compositae and Teasaceae. Asteraceae achenes develop from the lower paracarpous ovary, formed by two carpels, with a single ovule. The seed coat is greatly reduced. The achenes bear various appendages that aid in dispersal. In many species, a tuft of hairs develops, sitting directly on the top of the achene or on a special, sometimes very elongated spout. There are different opinions regarding the morphological essence of the crest.
Figure – Achenes of Asteraceae: 1 – dandelion; 2 – cornflower
Lysicarpous fruits. Lysicarpous fruits are characterized by a central column. The original type of lysicarpous fruit is a capsule, derived from a syncarpous capsule. In Cloveaceae, most genera have true lysicarpous capsules. Lysicarpous capsules are characterized by incomplete opening, not by valves, but mainly by denticles. The number of the latter may correspond to the number of carpels, or may be twice as large.
Single-seeded lysicarpous fruits (most often upper and dry) are often found. They are found in some carnation (Herniaria) and related families (amaranthaceae, goosefoot, vinaceae) and in the buckwheat order.
Infertility. Infertility is understood as the result of the fusion and transformation, as it were, into one fruit of several fruits arising from individual flowers of one inflorescence. For example, the so-called mulberry of a mulberry is an infructescence formed by fused fruits, the edible colored part of which, in fact, represents overgrown perianths. However, wineberries or figs are often considered as infructescence, although free, being in a hollow container, the fleshy walls of which are formed by the axes of the inflorescence. From this example it is clear that the concept of infertility is used in a broad sense.
Lecture: Fruits.
A fruit is a specific organ of flowering (angiosperm) plants, which is a closed container for seeds. It can develop without double fertilization and without seeds (with apomixis).
The variety of fruits is enormous, and they have long attracted the attention of researchers. Already in the 16th century, Caesalpini created an artificial system of classifying flowering plants based on fruit types. In the 18th century, the German scientist Gertner defined the science of fruits, the features of their structure, ontogenesis, ecology and distribution as carpology(lat.carpos– fruit).
People's close attention to the study of fruits is not accidental. Fruits are the basis of our life and have a huge meaning in human life:
1) food (starchy foods containing protein, fruits, drinks, vegetables, spicy foods);
2) feed (beans, vetch, oats, etc.);
3) oilseeds (tung, hemp, sunflower, etc.);
4) medicinal (hawthorn, raspberry, rose hip, etc.);
5) fibrous (cotton);
6) ornamental (bottle gourd, etc.).
The whole fruit or its parts (pericarp, seeds) can be used.
A fruit is the gynoecium of one flower modified after double fertilization or apomixis, sometimes with other parts of the flower growing to it.
Thus, a fruit is a flower modified after double fertilization or apomixis.
Fetal functions:
1) protection for seeds;
2) dissemination ( lat.disseminare– distribute) – the process of seed dispersal.
The meaning of fruits in nature:
1) ensure the settlement, reproduction and survival of plants (see fruit functions);
2) food for animals.
Fruit traits in plants are hereditarily stable. You can often identify a plant by its fruit. IN different groups Angiosperms, the evolution of fruits went in its own way, but precisely in the direction of strengthening their functions => some fruits became multi-seeded, others - few or single-seeded. But in any case, devices appeared that contributed to their better distribution.
There are cases when the same species, and even on the same plant, can produce fruits of different structures or having different physiological characteristics(for example, timing of germination). This phenomenon is called heterocarpy(diversity). It is found, for example, in calendula, in the families Asteraceae, Lamiaceae, borage, cruciferous, etc. (two dozen families are known). Heterocarpy has adaptive significance. Thanks to it, the distribution of fruits is improved (fruits different shapes spread differently), plants have backup dissemination pathways and plant survival improves (since different fruits germinate in different terms) => the adaptability of plants to environmental conditions improves.
Well-known carpologists in their circle worked at the Ulyanovsk Pedagogical University: Levina R.E. and Voitenko V.F., who studied heterocarpy.
The structure of the fruit.
The fruit consists of pericarp (pericarp ) and seeds. The pericarp usually has 3 layers: outer ( exocarp), average ( mesocarp) and internal ( endocarp). These layers can vary in consistency and structure or be of the same type. For example, in a cherry (the fruit is a drupe): the exocarp is filmy, the mesocarp is juicy, fleshy, and the endocarp is hard, stony and forms a stone enclosing the seed. The pericarp typically develops from the walls of the ovary, but sometimes other parts of the flower (bases of stamens, petals, receptacle, etc.) can also be involved in its formation.
Classifications of fruits.
Many attempts have been made to classify fruits, but due to the wide variety of fruits, no classification is ideal. All classifications of fruits can be combined into 2 groups:
1. Ecological-morphological (biological) - based on external characteristics (consistency of the pericarp, number of seeds, nature of opening, method of distribution).
2. Genetic (evolutionary) – taking into account the origin and evolution of fruits. These classifications are based on the type of gynoecium that forms the fetus.
1. Ecological and morphological classification of fruits.
Main classification characteristics:
1) character (consistency) of the pericarp (juicy or dry ). In dry fruits, all 3 layers of the pericarp are dry (membranous, fibrous, stony, leathery, etc.). Juicy fruits do not necessarily have all the layers juicy, one or two are enough.
2) number of seeds. There are: multi-seeded , low-seeded And single-seeded fruit.
3) nature of fruit opening. There are: a) drop-down multi-seeded fruits, b) disintegrating low-seeded fruits, c) non-tearing single-seeded fruits.
Decaying fruits are divided into two groups depending on how they disintegrate: along the fusion line of the carpels ( fractional fruits) or across ( articulated fruit). The part of the fractional fruit is called mericarp, and articulated - member. They usually consist of a closed portion of the pericarp with a single seed inside.
4) devices associated with the distribution of fruits (wings, crests, trailers, etc.).
Thus, in general view The classification of fruits is as follows.
Fruit |
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Dry | Juicy |
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Multi-seeded dehiscent | Low-seeded disintegrating | Single-seeded non-opening | multi-seeded | single-seeded |
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Leaflet Box | fractional | articulated | Nut Caryopsis | Berry | Monomeric | Polymer |
drupe | polydrupe Strawberry |
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Vislopordnik Diptera | Jointed bean Articulated pod |
1. Opening dry multi-seeded fruits.
A) Leaflets - a type of dry multi-seeded fruit that opens along the ventral suture.
Leaflets
Single-sheet multi-sheet
(formed from one carpel) (formed from several
apocarpous carpels
polymer gynoecium)
Leaflets are more common in primitive families and are usually formed from the apocarpous gynoecium. Found in Magnoliaceae, Ranunculaceae, some Rosaceae, etc.
B) Bean - a dry, usually multi-seeded fruit, formed from an apocarpous monomeric gynoecium and dehiscent along the ventral suture (the place of fusion of the carpels) and the dorsal vein (the midrib of the carpel). The seeds inside the bean are attached to the valves of the fruit. The bean is characteristic of the legume family. Some plants produce single-seeded beans - bobbies, and the ability to open is lost.
IN) Pod - a dry, usually multi-seeded fruit, formed from a paracarpous gynoecium, opening with two valves along the line of fusion of the carpels. Inside, the pod has a false septum (formed by placentas) to which sections of carpels with seeds are attached in the form of a frame. The pod is characteristic of the cruciferous family. When the seeds in a pod are reduced to several or one, the pod is shortened and called pod.
G) Box – a dry multi-seeded fruit of various structures and with different opening patterns. Often the boll is opened by valves (along the line of fusion of carpels or along the central veins) (cotton, iris), less often - by valves (for example, in poppy), or by a lid (in henbane, amaranth). The capsules are formed from the coenocarpous gynoecium.
2. Dry decaying fruits with few seeds.
Such fruits always contain a strictly constant, small number of seeds, with each seed isolated from the others by a part of the pericarp and often has special adaptations for spreading or germination.
A) Fractional fruits– disintegrate along the longitudinal line of fusion of the carpels onto the mericarp (“meros” Greek – part). Fractional fruits are formed from the coenocarpous gynoecium.
For example, the fruit of Umbrella - dwarf splits into two mericarps, which hang on a split stalk (carpophora).
In Lamiaceae and Borageaceae the fruit coenobium breaks up into 4 nut-shaped mericarps - erama(the gynoecium is formed by 2 carpels and the disintegration of the fruit occurs along and across the carpels).
Diptera maple tree splits into two mericarps, each of which has wing-like outgrowths (adaptations for gliding with the wind).
B) Jointed fruits– disintegrate across the line of fusion of the carpels along special dividing sutures-constrictions into segments.
Such fruits are found in legumes - a segmented bean (for example, kopeck, kopeck) and in cruciferous plants - a segmented pod (for example, wild radish).
3. Single-seeded, indehiscent fruits.
A) Nut – a single-seeded, indehiscent fruit with a woody pericarp (for example, hazel). Hazel has a nut enclosed in plus protective structure formed by overgrown bracts. The nut fruit is characteristic of the beech family, which includes beech, real (edible) chestnut, and oak. The oak fruit is a little specific, its pericarp is softer, leathery, this fruit is called acorn .
B) Achene - a single-seeded, indehiscent fruit with a leathery pericarp that does not adhere to the seed. Such a fruit, for example, in Compositae. Often the achene carries additional devices associated with propagation: crests, parachutes, trailers, etc.
IN) Caryopsis - a single-seeded, indehiscent fruit with a leathery pericarp attached to the seed. Characteristic of cereals, and may also have various adaptations for spreading (for example, feather grass).
4. Juicy multi-seeded fruits.
A) Berry – a juicy fruit with a fleshy pericarp and many seeds. The fruit is formed from a coenocarpous gynoecium (very rarely single-seeded or few-seeded). In the berry, the juicy endocarp and mesocarp merge with each other and the border between them is invisible, the exocarp is filmy. (for example: currants, gooseberries, tomatoes, blueberries, etc.).
B) Pumpkin - a fruit very similar to a berry, but differs in a leathery (sometimes woody) exocarp and highly developed juicy (sometimes fibrous) placentas. Characteristic of plants of the pumpkin family (cucumber, melon, pumpkin, watermelon). In watermelon, the endo- and mesocarp are homogeneous and the placentas are not so pronounced. Pumpkin is formed from a paracarpous gynoecium with an inferior ovary.
IN) Apple - a juicy multi-seeded fruit with a filmy exocarp, a juicy, fleshy mesocarp and a cartilaginous endocarp forming chambers containing the seeds. An apple is a specific type of fruit found in the apple subfamily of the Rosaceae family and is formed from a syncarpous gynoecium with an inferior ovary (apple, pear, rowan, hawthorn, etc.).
5. Juicy single-seeded fruits.
They can be divided into monomeric(or simple), formed from a single pistil (moreover, the gynoecium can be either apocarpous monomeric or coenocarpous) and polymer(prefabricated or compound), formed from several or many pestles.
A) Monomeric fruits include drupe - a juicy single-seeded fruit with a filmy exocarp, a fleshy juicy mesocarp and a hard stony endocarp, forming a stone containing the seed (cherry, plum, apricot, sea buckthorn, date).
B) Polymer (prefabricated) fruits include Polydrupe (raspberries, blackberries, drupes), consisting of individual small drupes. The fruit is formed from a single flower with an apocarpous polymeric gynoecium. The fruitlets are in close contact with each other and are attached to the convex receptacle that remains after the flower fades.
2. Genetic classification of fruits.
The first genetic classifications appeared at the end of the 19th century. One of the generally accepted classifications in our country is the classification of Rosa Efimovna Levina, which was proposed in the 60s of the 20th century.
This classification is based on type of gynoecium.
In accordance with this, the following types of fruits are distinguished: 1) apocarpic ( apocarpy); 2) coenocarpous ( coenocarps), which in turn are divided into syncarpies, paracarps And lysicarps.
Within these groups, fruits are divided according to the following characteristics: Apocarps are divided into: 1) polymeric and monomeric
2) single-seeded and multi-seeded
Each group of coenocap fruits is divided into: 1) upper and lower (according to the position of the ovary); 2) single-seeded and multi-seeded. Among syncarpous fruits, low-seeded fruits are also distinguished.
There are more small units: in each of the groups, juicy and dry fruits, dehiscent and non-dehiscent fruits, etc. are grouped separately.
The advantage of this classification is that it allows one to trace the main directions of fruit evolution. One of the main directions of fruit evolution is oligomerization. High quality oligomerization manifests itself in the fusion of carpels and the transition from apocarpous to coenocarpous fruits.
Quantitative oligomerization is manifested: a) in a reduction in the number of carpels to 1 (for example, from apocarpous polymeric fruits to apocarpous monomeric ones); b) reducing the number of seeds to 1 (from multi-seeded fruits to single-seeded ones).
This pattern can be seen most clearly in the example of apocarpous fruits.
1. Apocarpous fruits (aprocarps).
The most primitive fruits are considered to be apocarpous polymeric fruits, formed from apocapic polymeric gynoecium. The part of the polymer fruit is called fruit.
The original type of apocarpous fruit is considered spiral multi-leaf (magnolia, swimsuit), consisting of numerous multi-seeded leaflet fruits. The evolution of this type of fruit went in two directions: a) towards reducing the fruitlets to a small number ( cyclic multi-leaflet, e.g. peony, columbine) and then to one ( one-sheet , for example, consolidum field). In this case, the apocarpous polymeric gynoecium becomes monomeric. From a single leaflet could be formed bean , when the nature of the opening changed, and the fetus began to open not only along the abdominal suture, but also along the dorsal vein. Through a segmented bean, a transition to a single-seeded bean (pod) could occur, when only one segment remains.
b) The second direction of evolution is a reduction in the number of seeds in fruitlets to 1 and the transition from a spiral multi-leaflet to multi-nut (fruit-nuts do not open, but fall one by one from the convex receptacle) (buttercup, adonis, cinquefoil, etc.). Among the apocarpous fruits there are various varieties of polynut: strawberry – the nuts are partially immersed in a fleshy, juicy, overgrown receptacle (strawberry), cynorhodium – the nuts are located inside the overgrown fleshy hypanthium (rose hips), lotus fruit, etc.. Associated with the polynut in its origin Polydrupe : the walls of the pericarp have become juicy. Then, as a result of further oligomerization, a transition to monodrupe (reduction in the number of fruitlets to 1) occurred (transition from apocarpous polymeric to apocarpous monomeric fruits). Juicy drupes are found in cherries, apricots, plums and some other Rosaceae; dry drupes are found in almonds.
2. Coenocarpous fruits.
They are divided into: syncarpous, apocarpous and lysicarpous, evolutionarily interrelated. According to the classification of R.E. Levina, syncarpous fruits evolved from apocarpous fruits, and from those fruits, paracarpous on the one hand, and syncarpous on the other.
Coenocarpous fruits evolve from polyspermous, formed by several carpels, to single-seeded fruits, in which the number of carpels, nests and ovules decreases to 1.
In each of the groups of coenocarpous fruits the following are repeated: boxes (upper and lower) – syncarpous – nightshade, iris; paracarpaceae – violet, orchids; lysicarpous - cloves; berries (upper and lower) – syncarpous – lily of the valley, blueberry; paracarpous – gooseberries, currants; lysicarpous - mistletoe, as well as drupes , dry and juicy (sea buckthorn, etc.). Each group also has its own specific types of fruits.
2.1. Syncarpous fruits.
Upper multi-seeded: syncarpous polyfoliate (cableweed), boll (henbane), berry (lily of the valley). Specific citrus fruit - orange (hesperidium) -juicy citrus pulp is formed by hair cells of the inner layer of the endocarp, filled with cell juice.
Upper low-seeded: fractional fruits – coenobium (Borage, Lamiaceae), fractional boll (mallow) and lionfish (maple).
Upper single-seeded: lionfish (elm, ash), nut (Linden).
Lower polyspermous: capsule (iris), berry (blueberry, banana), specific fruit - apple.
Lower low-seeded: fractional fruit of Umbelliferae dwarf And double drupe Merenaceae
Lower single-seeded: lionfish (birch), nut (hazel), acorn (oak), dry drupe (walnut).
2.2.Paracarpous fruits.
These fruits are less diverse than syncarpous ones.
Upper multi-seeded: capsule (violet, poplar), berry (papaya), specific fruits: pod And pod (cruciferous) (evolution went from a pod through a segmented pod to a single-seeded pod).
Upper single-seeded: grain (cereals), nut-shaped pod (sverbiga), dry drupe (coconut).
Lower polyspermous: capsule (orchids), berry (gooseberry, currant), specific fruit – pumpkin (cucumber, pumpkin).
Lower single-seeded: achene (Asteraceae), drupe (Lox).
2.3. Lysicarpous fruits.
Least diverse, rare. Almost all of them are top. The most typical are multi-seeded capsules with a column (cloves, primroses) and single-seeded achenes (buckwheat, amaranth, goosefoot).
Infertility.
In a narrow sense: infructescence is a collection of closely adjacent and often fused fruits.
In a broad sense: infructescence is a collection of ripe fruits of one inflorescence.
Examples of fruits: a pineapple - all the pistils are fused together, the axis of the inflorescence fuses with the ovaries and bases of the covering leaves into one juicy, fleshy fruit.
Figs form fruit - synconium . The axis of the inflorescence is pitcher-shaped, concave, fleshy. Inside, first there are numerous small flowers, and then fruits - nuts immersed in a fleshy, overgrown axis.
Glomerulus beet consists of several fused fruits. Therefore, during germination, several seedlings are formed. Nowadays, beet varieties have been developed whose ball contains 1 seed (“Single-sprouted”).
Infructescence is also formed in cattail (cob), strawberry clover (head), etc.