Pharmaceutical Botany for International Students

PHARMACEUTICAL BOTANY

FOR INTERNATIONAL

STUDENTS

Minsk BSMU 2017

МИНИСТЕРСТВО ЗДРАВООХРАНЕНИЯ РЕСПУБЛИКИ БЕЛАРУСЬ БЕЛОРУССКИЙ ГОСУДАРСТВЕННЫЙ МЕДИЦИНСКИЙ УНИВЕРСИТЕТ КАФЕДРА ОРГАНИЗАЦИИ ФАРМАЦИИ

ФАРМАЦЕВТИЧЕСКАЯ БОТАНИКА ДЛЯ ИНОСТРАННЫХ СТУДЕНТОВ

PHARMACEUTICAL BOTANY FOR INTERNATIONAL STUDENTS

Учебно-методическое пособие

Минск БГМУ 2017

2 УДК 615.1:581(075.8)-054.6 ББК 52.81(81.2 Англ-923) Ф24

Рекомендовано Научно-методическим советом университета в качестве учебно-методического пособия 21.12.2016 г., протокол № 4

А в т о р ы: проф. Н. С. Гурина; доц. О. А. Кузнецова; доц. О. В. Мушкина; ст. преп. М. В. Волочник

Р е ц е н з е н т ы: канд. с.-х. наук, доц., зав. каф. ботаники Белорусского государственного университета В. Д. Поликсенова; канд. мед. наук, доц. каф. биологии Белорусского государственного медицинского университета Л. М. Сычик

Фармацевтическая ботаника для иностранных студентов = Pharmaceutical Ф24 botany for international students : учеб.-метод. пособие / Н. С. Гурина [и др.]. – Минск : БГМУ, 2017. – 112 с. ISBN 978-985-567-648-6. Содержит теоретический материал по фармацевтической ботанике, схемы, рисунки и представ- ляет собой краткий курс лекций. Предназначено для иностранных студентов 1–2-го курсов, обучающихся на английском языке.

УДК 615.1:581(075.8)-054.6 ББК 52.81(81.2 Англ-923)

ISBN 978-985-567-648-6 © УО «Белорусский государственный медицинский университет», 2017

3 LECTURE № 1 BOTANY AS A GENERAL DISCIPLINE. BASIS OF CLASSIFICATION OF LIVING ORGANISMS. ALGAE

Pharmaceutical Botany is a general discipline, which is forming theoretical knowledge and practical skills which are necessary in a future pharmaceutist’s working. Botany, as a science, studies external and internal plant’s structure, features of processes of vital activities, classification, intercommunication with environment conditions, spread in nature, meaning in the nature and human life. Complex study of the plants caused a necessity in selection from Botany some fundamental disciplines: Morphology of plants, Anatomy, Physiology, Embryology, Systematic, Geography and Geobotany, Ecology of plants, Paleobotany. Meaning of Botany in Pharmacy: 1. Anatomy of plants as a basis of microscopical analysis of medicinal plant raw material. 2. Morphology as a basis of macroscopical analisys of medicinal plant raw material. 3. Physiology of plants is necessary for studying the metabolism and the accumulation of biologically active substances. 4. Systematic of plants is necessary for identification of plants in nature. 5. Botanical Geography as a basis of rational storing of medicinal herbs. 6. Ecology of plants (phytocoenology) for the introduction of plants into the culture. Botany as a science appeared under the influence of practical human needs. It developed along with human society. First scientific knowledge about the plants is found in the works of Greek classics in the 4th–3rd centuries BC — Aristotle and Theophrastus considered as a ―father‖ of Botany as he was the first to classify plants into trees, shrubs, subshrubs, herbs (perennial, biennial, and annual). Plant is a living organism. Every vegetable organism is an open, autoregulated, self-replicated system, which is characterized by streams of matter, energy and information. The stream of matter lies in the basis of metabolism. Metabolism is a totality of assimilation and dissimilation reactions, known as anabolism and catabolism. Vegetable metabolism may be subdivided in primary and secondary ones. Primary metabolism includes synthesis and disintegration of proteins, carbohydrates, fats, nucleic acids, ascorbic acid; secondary metabolism includes formation and transformation of organic compounds of other chemical classes (alkaloids, tannins, glycosides and others). The stream of energy is connected uninterruptedly with the stream of matter and is realized through synthesis and disintegration of ATP, which occurs on assimilation and dissimilation accordingly.

4 Organism is an open system, as it cannot exist without external energy supply for most of the plants the source of energy is the Sun, and this energy is stored in the form of chemical bonds in the process of photosynthesis. Such organisms are called autotrophs (phototrophs). Unlike the plants, animals, fungi, and bacteria use for their vital activities energy, which frees on disintegration of organic substances (these substances are absorbed by these organisms). The organisms are called heterotrophs. The stream of information lies in the basis of self-replication of organisms and is connected with nucleic acids functions. Plant, as an integrated organism, has all the characteristic biological levels of organization on the le vels of molecule, cell, tissue, organ, ontogene, population, species, ontogenesis, and biosphere.

THE BASIS OF LIVING ORGANISMS CLASSIFICATION. ALGAE Systematics is a science which studies the variety of all existing and extincting organisms. Scientists consider that about 500 thousand species of plants and about 2 million species of other living organisms exist on the Earth at the present time. The purpose of Systematics is revelation, description and classification of organisms into the system. Divisions of Systematics are Taxonomy, Nomenclature and Phylogenetics. Taxonomy represents theory and practice of organisms’ classification, i. e. distribution of all the present known organisms by a certain system of a subdominant categories on the basis of their similarities and distinctions. Similarities and distinctions are determinated with the help of different methods. They are estimated by specially developed criteria. Nomenclature is a totality of all existing names of taxons. Phylogenetics studies historical development of the world of living organisms (Phylogeny) and congeniality of organisms during this development. Basic terms of Systematics are is taxons and taxonimical categories. Taxonomical categories desgribe certain levels in the hierarchic classification. Taxon is a certain group of organisms, existing or extinct. The basic taxon, which unites species most connatural to each other — is a specie (species). Plants, which are used by a human, must be determined until specie, sometimes even more exactly. The first determination of specie was given by C. Linnaeus. Linnaeus considered species permanent and changeless. There is no one and generally accepted determination, which is equally suitable for plants, amphimicticing, and plants, agamogeneticing or reproducing vegetative, of a specie until the present moment. We will introduce one example from the modern determinations.

5 Specie is a group of units similar by morphological, physiological, biochemical and other features randomly mating with each other, giving a fertile offspring and living on the certain territory called areal. There are varieties inside the specie, subspecie, which can give new species in the process of evolution. There is a continuous raw of forms within the specie, no matter how big the intraspecies variability is. At the same time discretisation exists even between the closest species. There are no transitional forms. Species have distinctions not only in external features, but in living conditions too. There are barriers for crossing. Genuses can be monotypic, oligotypic and polytypic. They can be divided into sections and subgenuses. Species unites into the genuses. Genus (genus) is the highest taxonomic category, uniting groups of relative species. For all taxons, beginning from genus, the scientific names exist. The names consist of one Latin word (uninominal). Binominal names, consisting of two Latin words are accepted for species: the first word is a name of genus, which belongs to the specie; the second word is a specific epithet, e. g. plantago major, or broadleaf plantain; urtica dioica etc. This binary nomenclature was introduced by C. Linnaeus in 1753. Taxonomical categories of a higher rank are as below: Family (familia) includes one or two genuses. The name of the family derivesfrom the name of genus with the inflexion ―aceae‖, for example: Convallaria – Convallariaceae. Order (ordo) is a systematic category, which includes several families. It is called with the inflexion ―ales‖. Class (classis) is a higher taxonomical category. The number of classes is not big. For example, angiosperms have two classes, Monocotyledones and Dicotyledones. The inflexion ―psida‖ is accepted for classes, but names that were used long ago are also acceptable. For example, Dicotyledones can be called Magnoliopsida or Dicotyledones. The biggest taxonomical unit in The Kingdom of Plantae is division (division). Divisions are called with an inflexion –phyta and distinguish from each other by fundamental features. They are stated in the order of an evolutional complicacy.

MODERN SYSTEM OF THE LIVING NATURE All the variety of organic world is divided into two empires: The Empire of Noncellular life, i. e. Noncellulata The Kingdom of Viruses, i. e. Virae The Empire of Cellular life, i. e. Cellulata The Domain of Procaryote, i. e. Procaryota The Domain of Eucaryote, i. e. Eucaryota 6 The differences between representatives of the domains Procaryote and Eucaryote: Procaryote Eucaryote 1. Cell size Under 10 µm 10–100 µm 2. Formalized nucleus Nucleoid Exists 3. DNA In nucleoplasm In nucleus 4. Mitochondria, plastids, Goldgi complex and etc. None Exists 5. Ciliae (if exist) Simple structure Complex structure (from flagellin) (undolipodium) 6. The cell membrane consists of: Mureine Cellulose or chitin 7. Proteins Without histones With histones 8. Movement of cytoplasm None Exists 9. Cell division Amitosis Mitosis, miosis 10. Amphimixis or syngamy None Exists 11. An ability of fixing the Nitrogen Exists None

Procaryote includes only microscopically small organisms, its cells don’t have a formalized nucleus. Procaryote is represented by one-celled or colonial organisms. Their cells are small, under 10 µm, i. e. one order less the cells of Eucaryote. Many of Procaryote have ciliae with a simple structure from protein flagellin. Glycopeptide mureine are often present in their membrane (to compare, cellulose is present in plant’s membranes, chitin in fungi’s). Their cells possess no mitochondria, chloroplasts (or green plastids), Goldgi complex. Ribosomes are smaller the Eucaryote’s ribosomes and they are distinguished in protein’s composition. The analogue of nucleus is genophore, which consists of DNA, proteins and RNA. Genetical system in the shape of circle is fixed on the cell membrane and corresponds to a primitive chromosome. Mitosis and miosis are absent, amphimixis or syngamy are very rare. Reproduction happens by a simple binary division of cell into two cells. Duplication of DNA happens before division. Bud reproduction appears sometimes. Process of delivery of hereditary information from one strain to another with the help of DNA pieces exists. Procaryote stands in a marked contrast from Eucaryote in a peculiarities of metabolism. Many of them are capable of fixing the Nitrogen from atmosphere. There is also Anaerobic Procaryote. Procaryote is the first organism that appeared on the Earth about 3.2–3.3 billion years ago. They include two kingdoms: Archaebacteria (Archaebacteria) and Eubacteria. The Kingdom of Arhaebacteria. Archaeobacteria. They distinguish from Eubacteria by a number of physiological and biochemical features. Cell membrane does not consist of mureine, but polysaccharides, RNA composition is very different from all the others oragnisms. Methane-producing bacteria, reducing CO2 to CH4, are the best known. They inhabit strictly anaerobic conditions. They can be used for biogas production. 7 The Kingdom of Eubacteria. Eubacteriabionta. They are the subject of microbiology studies. With the help of colouration method by aniline colours, which was suggested by H. C. Gram, bacteria are divided into gram-negative microorganisms (The Subkingdom of Oxyphytobacteria, Anoxyphytobacteria, Schizomycetes, Spirochaetobacteria) and gram-positive microorganisms (The Subkingdom of Actinobacteria, Bacteriobionta, Micoplasma). The name comes from Greek ―bacterion‖ which means ―rod bacterium‖. There are a number of morphologic groups based on the shape of cells: coccus (roundshape), bacilli (rod bacterium), vibrio (arcuated), spirilla (gyrose). They often possess ciliae and villi for moving by slipping. Gram-negative microorganisms have thick-walled shell which does not get a colouration by Gram. The main compound of the membrane is mureine. The cell of many of bacteria is surrounded by a layer of mucus. Storage compound is a volutin, sometimes amylum or glycogen. They do not form sporules. Gram-positive bacteria are capable of forming endospores. Membrane of endospore is very stable; it can survive very high and low temperatures, dehydration and other unfavorable conditions, saving the protoplast for a long time. In favorable conditions (warm, humid, breeding ground) endospores sprout. There are a great number of such microspores in the atmosphere, water, ground and practically everywhere. Autotrophic and Heterotrophic bacteria are known. In their turn, Autotrophic bacteria are divided into chemotrophic bacteria and phototrophic bacteria. Chemotrophic bacteria can live without the light and organic substances upon condition of presence CO2, O2 or other inorganic compounding capable of oxygenation. There is only a little number of such bacteria, e. g.: nitrobacteria, sulfur bacteria, iron bacteria. Their role in the substance circulation (biogeochemical cycles) is great. Many of the most important reactions, e. g. nitrogen bonding, and nitrification can only be proceeded with the participation of bacteria. Phototrophic bacteria contain bacteriochlorophyll and they are capable of photosynthesis, but without excreting oxygen. They inhabit in a shallow clear water reservoir. Heterotrophic bacteria use organic substances for getting energy. Saprophytes, parasites, sometimes symbiotes are among them. The main role of bacteria in the nature is mineralizing of organic compounds. The activity of bacteria is used by man for getting many alimentary and technical products — kefir, cheese, koumiss, different ferments, alcohol, citric acid, antibiotics and others. The bacteria are also used in fermentation of vegetables, in genetic engineering. The negative role of bacteria is connected with ration spoilage, wood spoiling, as well as with causing plant’s animal’s and humans’ diseases.

8 The Domain of Eucaryote: Eucaryota. It is divided into several Kingdoms: Protoctista, Animalia, Fungi (Mycota), Plantae (Vegetabilia). The Kingdom of Protoctista. Eukaryotic organisms, inhabiting water reservoirs, one-celled or colonial, with a furcated thallome hve no differentiation on organs. Embryonic stages are absent. Nutrition is holozoic (by swallowing), osmotic (by soaking), and autotrophic. They include: The simplest animal organisms — Protozoa Fungaceous protoctista — Divisions: – Oomycota (Phytiphtora — affects top and tubers of potato, Plazmopara — affects grapevine); – Chytridiomycota (Synchitrium endobioticum — cancer of potato, Olpidium brassicae — ―a black stipe‖ of cole (cabbage)); – Myxomycota — Plazmodiophoromycota. Protoctista — Algae. Common characteristics: 1. Their body never divides into organs and is presented by thallome. Sometimes they have one-celled outgrowths called rhizoid. 2. They inhabit water reservoirs, ground, stones, cliffs, trunks, can be symbionts of Lichenes. 3. Their cells are built like plants cells. They have a polysaccharide membrane (some of them have a pellicle). Plastids (chromatophore) are different shaped with pyrenoids. 4. A variety of pigments, e. g. chlorophylls, caratenoids, phycobillins (fucoxanthin, phycoerythrin, phycocyanin and others)

CLASSIFICATION The Division of Euglenids The Division of Red Algae (Rhodophyta) Red algae are alike with blue-green algae by a number of pigments (chlorophyll a, d, phycocyanin, phycoerythrin) and distinguish from all other plants by this. Their storage compound is a special red algae’s amylum. Their cell membrane contains special pectic substances, which are used by a human under the name agar-agar in microbiology and confectionary industry. Body of Red algae — thallome is in a form of pluricellular fibres, forming pseudaparenchymic pallets (laminas). They attach the substrate by rhizoides. The most deep dwelling inhabitants of seas. Their reproduction can be vegetative, sexual, and asexual. The most important characteristic of their development cycle is an absence of flagellate stages; spores and gametal cells are always motionless, they move with the help of water run.

9 The typical representatives are Porphyra, Nemalion, Callithamnion. We will view sexual reproduction of Red algae at the example of Nemalion, which inhabit the Black sea. Thallome of this alga consist of thin fibers, fastened into batch. Oogonium (sporophydim) is bottle-shaped and called carpogonium. An ovum grows up in a wide part of oogonium called belly. An upper side of carpogonium is called trichogyne. Motionless male gametes spermatiums, grow up in a numerous antheridium. They passively move with water run, adhere to trichogyne, protoplasts of spermacium and ovum merge. A carpospores forms from appeared zygote and it gives a beginning to a new organism. The asexual reproduction is performed by tetraspores.

А 1 2 B 3 4 Figure 1. A — Batrachospermum: 1 — outward appearance of thallomee, 2 — part of thallome. B — Nemalion: 3 — cross section of thallome, 4 — carpogonial branch

The Division of Diatoms: Baccilariophyta. They stand in marked contrast from all other groups of Algae by a presence of a hard siliceous cover (a frustule). One-celled or colonial species. Cellulose membrane is abscent. A frustule consist of two parts — epitheca and hypotheca. Chloroplasts are in the shape of grains or lamellas. Pigments — chlorophyll, carotin, xantophyll, diatomin. Their storage compound is fatty oil. Reproduction is vegetative and sexual. They live practically everywhere in seas and fresh water reservoirs. Their representative is Pinnularia. The Division of Brown algae — Phaeophyta. They are pluricellular inhabitants of seas, the biggest of all the known algae, sometimes 60 meters in length. Their cells have nucleus, one or two vacuoles, their covers are very thin. Chloroplasts are coloured in a brown colour (pigments: chlorophyll a, c, carotin (carotene), xantophyll, fucoxanthin). A storage compound is laminarin, mannitol and fats. Reproduction is vegetative, sexual and asexual with a strict alternation of generations by isomorphic or heteromorphic type.

Figure 2. The Diatoms: 1 — Pinnularia; 2 — Navicula; 3 — Synedra; 4 — Pleurosigma; 5 — Melosira; 6 — Gomphonema Representatives — Laminaria, Fucus.

Figure 3. Brown Algae: A — Fucus, B — Nereotsistsis, C — Nereocystis, D — Macrocystis, E — Laminaria japonica (―sea cabbage‖)

The Division of Green algae — Chlorophyta. The biggest division among algae includes about 5 thousands species. Its representatives are very speciose by a external appearance: one-celled, pluricellular, siphonal, filamentous and placoid algae. They inhabit in a fresh or sea water, also on the ground. The specific feature of Green algae is that their kit of pigments is practically like a higher plants kit (chlorophyll a and b, carotinoids). Chloroplasts have a two-membranous cover, speciose by a shape, pirenoids can be present. Cell-memrane consists of cellulose, pectic substances. There are motile forms with undolipodium. Storage compound is amylum, rarely it is oil. 11 Representatives: Chlamydomonas — one-celled alga, reproductive process is isogamic. Spirogyra — filamentous algae. Reproductive process occurs by conjugation. Caulerpa is a non-cellular structure (siphonal), outwardly reminds pedicellate plants. It represents a giant cell with outgrowths 50 cm in legth sometimes, having one protoplast with an endless vacuole and numerous nucleuses.

A Figure 4. Green algae. From left to right. Caulerpa. Spirogyra, Ulva: A — an external appearance of thallome; B — transverse incision of thallome

A Division of Charophytes is Charophyta. Being pluricellular, their body is divided into parts. The external appearance is similar to external appearance of higher plants. Reproduction vegetative and sexual (oogamy). Oogamy has a characteristic structure, with a cover which consists of twisted cells, formatting a crown on the top. Antheridium is round-shaped. Zygote after a dormant period grows into a new plant. A representative is Chara fragilis.

Figure 5. Charophytes. Left to right. Chara: external appearance of thallome, transverse incision of thallome; Nitella 12 The meaning of Algae: a huge role of creating the organic substances and oxygen on the planet, in cycle of matter, in nutrition of water reservoirs’ inhabitants. Algae can perform self-purification of waters. Many of algae are indicators of habitat areas pollution. They may be used as food for human and live-stock animals, and in quality of fertilizers. Algae are used for getting agar- agar (Anfelcia), sodium alginate (glue) for producing soluble fibers that are used in surgery. Laminaria, Fucus, Spirulina are used in a medicine. ―Sea cabbage‖ is a mix of Laminaria and Porphyra.

13 LECTURE № 2 THE KINGDOM OF FUNGI/MYCOTA/MYCETALIA. COMMON CHARACTERISTIC, CLASSIFICATION, REPRESENTATIVES

Fungi are heterotrophic organisms, separated into individual Kingdom. Common characteristics: 1. Fungi are saprophytes, symbionts, parasites. 2. Their body is represented by mycelium, or mushroom spawn, consisted of thin branched mycelial filaments (hypha) with an apical growth and lateral branching. Mycelial filaments may be haploid, dikaryotic (only cytoplasms fuse and nuclei of cells exist independently), diploid. 3. Tissue created by mycelial filaments is called plectenchyma (false tissue). 4. Fungi cell has a chitinous cover (yeast fungi’s cover consists of glucosanes). There are ribosome, mitochondria and nuclei in the protoplast. Glycogen is stored in vacuoles. The product of a protein exchange is urea. 5. Reproduction may be vegetative, sexual and asexual. Vegetative reproduction occurs by parts of mycelium, asexual occurs by means of formation of special spores (they are called conidia), sexual reproduction occurs by means of fusion of gametes (gametangiogamy) or fusion of somatic cells (somatogamy). Maturation of male and female reproductive organs (archegonium and antheridium) and a following fertilization happens inside mycothallus which is formed by mycelium. Spores of sexual reproduction (basidiospore, ascospore) develop from the formed zygote (diploid or dikaryotic). Classification hasn’t been determinate yet. Nowadays 5 divisions are separated: 1. Zygomycota 2. Ascomycota 3. Basidiomycota 4. Deuteromycota — Fungi imperfecti 5. Lichenes, Phycomycota The Division of Zygomycota have a well-developed non-cellular mycelium; sporangiospores are motionless; reproductive process is zygogamy. Representative is a mould called Mucor Mucedo. It develops as a saprophyte on vegetable humid products. Mycelium after germination of spore develops gross, penetrating substrate in all directions. Sporangiophore, ending with a round-shaped sporangium, ascend vertically from mycelium. Spores after opening the sporangium spread with air flows. Spores germinate in a new mycelium in favorable conditions. This is an asexual reproduction. Sexual reproduction happens by means of mycelial filaments from different individuals. Cells (gametagangium) on the endings of mycelium divide, fuse, and a zygospore with a diploid nucleus is formed. It grows, and is covered

14 with a hard spiny cover. Unbranched hypha with embryonic sporangium above is formed from zygospore after germination. Before forming spores, a reduction division happens. Haploid spores give heterosexual haploid mycelium. Ascomycota or sac fungi is one of the biggest divisions of fungi. Approximately 30 thousand species are known which is 30 % of all known species of fungi. The main feature of ascomycota is forming sacs or ascus, containing fixed number of ascospores (usually 8) as a result of reproduction process. Mycelium is pluricellular consisting of polykaric or mononuclear cells. Male and female reproductive organs develop on a Pluricellular mycelium. Female reproductive organ (archicarp) consists of ascogone and trichogyne (dilated and narrow part); male reproductive organ consists of one cylindrical cell (antheridium). Ending of antheridium grows together with trichogyne and its contents transfuse into archicarp. However, nuclei are not fused, but approach close to each other forming a dykaryonic zygote, whose germinating gives dykaryotic mycelium. Outgrowths or ascogenous hyphas develop on it. Sacs or ascus are formed on their endings by fusing nuclei. Then 3 divisions of nucleus happen, one of them is a reduction. In each sac 8 ascospores are formed. They give a haploid mycelium after germination. Therefore, in cycle of ascomycota 3 stages are interchanged: – Long haploid stage, during which asexual reproduction happens. – Short dikaryonic stage (dikaryonic mycelium, ascogenous hyphas). – A very short diploid stage (young sac with a diploid nucleus). In a cycle of ascomycota development an asexual reproduction plays a huge role. Spores of an asexual reproduction — conidium — are formed on haploid mycelium on conidiophores. Conidium develop in a period of fungi vegetation and serves for their mass spreading. Sacs of most ascomycota are alongated, they can have vellus and they can be assidenous. Layer of sacs is called hymenium. Sterile filaments or fibres paraphysis appear between sacs. Apparently, they contribute to spreading of ascospores. During growth and branching ascogenous hyphas are overgrown by usual haploid hyphas from all sides, and form a tight entanglement — a fruiting body (Ascocarps). Three types of a fruiting body are characteristic for sac fungi: 1. Cleistothecium (Cleistocarp) — round, completely isolated fruiting bodies. Sacs free from them only in the process of destruction of a cover. 2. Perithecium (Pyrenocarp) — half-isolated frit bodies, round or pyxidated with a narrow hple on the top. 3. Apothecium — widely opened during maturation fruiting bodies. They are usually patelliform, discoidal or round-shaped. On their upside hymenium is situated.

1 2 3 Figure 6. Types of a fruiting body sac fungi: 1 — Cleistothecium, 2 — Perithecium, 3 — Apothecium

According to a presence of fruiting body ascomycota are divided into 2 subclasses: Hemyascomycetidae (exoascale fungi) and Euascomycetidae, Loculoascomycetidae. All the primitive ascomycota belong to a small subclass of hemyascomycetidae. Fruiting body are absent and their sacs develop alone or in layer directly on their mycelium. The most common representatives are yeast fungus. They do not form typical mycelium and their vegetative cells bud or divide. Ascospores are formed inside sacs represented by single cells. Body of yeast fungus in the simplest form represents one cell, which has all general structures, characteristic for fungi. Sometimes cells may unite into different more or less strong structures in form of false or real mycelium. Sizes of single yeast fungus’ cells are 1–10 µ. The Form of yeast fungus’ cell depends on the way of vegetative reproduction. It may happen by budding or division. Some species of yeast fungus have a heterogamic reproduction process. Yeast fungus include very big and various group of organisms. Genus Saccaromyces has the greatest meaning for a human. This genus unites both natural and productional species. Their most common feature is the capacity for an active attenuation of sugars with a production of maximum, comparing to other yeast fungus, amount of alcohol. Beer, wine, baker’s yeasts belong here. The system of Euascomycetidae is generally based on the structure of fruiting bodies. As a matter of convenience they are divided into groups of orders accordingto a type of fruit body and sacs. A group of orders is Flask fungi (Pyrenomycetes). The fruiting bodies are pyrenocarps, less often cleistocarps. Outstanding interest in this group represents The Order of Claviceps, represented by Claviceps purpurea, which parasitize on the grass family (cereals). Cycle of Claviceps’ development. In the end of summer dark-violet, slightly curved, long horns (sclerotium) appears on rye’s heads. They consist of a tight entanglement of hyphas, which are filled with nutrient materials and lost the water. Sclerotium winter, falling on the ground and red heads on legs germinate from sclerotium in the spring. The red heads on legs are called 16 stroma. Stroma is a few fruiting bodies. Ascogen and antheridium develop in each stroma. Fertilization happens and a dikaryonic zygote is formed, ascogenous hyphas with dikaryons are formed from the zygote. Before forming ascospores the fusing of nuclei and 3 serial divisions happen. As a result, 8 haploid ascospores are contained in each sac. Being thrown from sacs ascospores get on heads’ flowers with the help of wind or insects. Then they germinate in a mycelium, which invades into pistil’s ovary. Mycelium oozes sweet sticky liquid, which is called ―honeydew‖ attracting insects. Mycelium breaks through the walls of ovary and develops a lot of spores — conidium, they are stuck to insects, together with a sweet liquid, and transferred to healthy flowers. By the end of summer mycelium is ruined by itself, ovaries start to sclerose and form sclerotium again. In such a way, in a cycle of Claviceps’ development typical for sac fungi stages: – long haploid, during which the asexual reproduction; – short dikaryonic (dikaryonic mycelium, ascogenous hyphas); – very short diploid (young sac with a diploid nucleus) are traced. Sclerotium of Claviceps contain alkaloids: ergotamine, ergometrine, ergokryptine and others.

Figure 7. The life cycle of Claviceps

The Order of Plectascales. Dermatophytes-fungi inhabit hairs, nails, and skin, causing dermatomycosis. A representative is Trichophyton. They have isolated fruit body — cleistocarps. Another representative is Emericellopsis terricola which produces cephalosporin.

17 The Order of Discomycete. Fruiting bodies — apotheciums. Representatives from genuses Gyromitra (saddle fungus) and Morchella are the most common. By an external appearance the fruit body represent mushroom (blewits), but its surface is irregular with cup-shaped extensions, the hymenium (a layer of sacs with sterile fibers — paraphysis) is situated on it. They are considered as potentially pathogenic fungi. The Order of Basidium fungi — Basidiomycetes. It is higher fungi with a pluricellular mucelium. Known are approximately 30 thousand species. They are distinguished from sac fungi by an absence of reproductive organs. Reproductive process is performed by fusion of two vegetative cells of a haploid mycelium, germinating from basidiospore. Fusion of cytoplasm happens, but nuclei forms dikaryons which then divide synchronously. Such dikaryonic mycelium, penetrating a substrate, may exist for a long time. Outgrowths — basidiums, are formed on the endings of dikaryonic hyphas, 3–4 basidiospores are formed on a basidium. Most of basidiomycetes form basidium with basidiospores on fruiting bodies, composed of dikaryonic mycelium. Basidium and paraphysis compose a hymenial layer; on fruiting body it is called a hymenophore.

A B C Figure 8. A — a scheme of sac’s developing; B — scheme of basidium’s developing: 1 — spores; C — Fruiting bodies of Basidium fungi

Sacs of sac fungi and basidium are distinguished by that that spores in sacs develop endogenous and basidiospores develop exogenous on the surface of basidium on thin outgrowths. Basidium of most basidiomycetes stays one-celled and called holobasidium, but some of them have a division and their basidium consists of four cells and are called fragmobasidium. Three subclasses: 1. Holobasidiomycetidae. 2. Heterobasidiomycetidae. 3. Teliosporomycetidae.

Figure 9. The life cycle of Basidiomycetes

Holobasidiomycetidae include two groups of orders: Hymenomycetes and Gasteromycetes. Hymenomycetes include 2 big orders: Polyporales and Agaricales (gilled mushrooms). Polyporales (polypores). Their representative is Fomes fomentarius (tinder fungus, false tinder fungus, hoof fungus, tinder conk, tinder polypore or ice man fungus). It has annual or perennial fruit body, hoof-shaped or snail- shaped; its consistence is juicy or dry, ligneous (woody). Mycelium is internal, developing in a living or dead wood, also in a forest soils, where there is mycorrhizal. Its hymenophore is boletus (tubicolous). It plants on an adult birch, continuing to develop on dead trees. It should be distinguished from fungus Inonotus obliquus (or chaga fungus), which develops on birch tree, rarely on alder, nut-tree (hazel). It has grumous fruit body in the shape of outgrowths and it is used in a quality of medical plant raw material. According to the State Pharmacopoeia of the Republic of Belarus, it can be supplied in the course of the year; it has a form of different-shaped pieces with a size under 10 cm, with a black badly cracked exterior layer. Another representative of the Family is Serpula domestica, a dangerous destroyer of domestic wood materials. It invades into a damp wood, causing its destruction. The Order of Agaricales (gilled mushrooms). They are fungi with a tabulate hymenophore, the fruit body has a shape of cap sitting on a stipe. On the downside of cap, numerous vertical directed pallets separate, hymenium develops on this pallets. Most of edible mushrooms relate to this family: 19 champignons, honey mushroom, yellow swamp brittlegill, wooly lactarius, ugly milk-cap, orange milky cap and etc. Some poisonous fungi are sulphur tuft, fly agaric, death-cup, etc. The Subclass of Teliosporomycota includes orders Ustilaginales and Uredinales. Some representatives are rusts and smut fungi. They don’t have a fruit body. Smut fungi consist of some hundreds of species and they are parasites. Mycelium is articulate, very branched, develop in intercellular space and invade by its haustorium into lumen. Flowers suffer most of all, because they can be totally destructed by mycelium of smut fungi. A huge number of smut fungi’s spores appear in a head or panicle of gramineous instead of grain harvest. Common bunt, loose smut and others are spread. All of them cause a huge damage to the nation’s economy. Rusts parasitize on cereals too. Brown and stripe rusts, affecting wheat, are spread, as well as other species of rusts that affect sunflower, lint and other plants. Deyteromycota — Fungi imperfecti. Together with Ascomycota and Basidiomycota, Fungi imperfecti represent one of the biggest suborders of fungi, which is about 30 % of all the known species. They are characterized by a Pluricellular mycelium, but they have only asexual reproduction — by conidium. Fungi imperfecti compose a group of the most dangerous parasites and saprophytes. Three orders are separated: 1. Hyphomycete. They are spread in the soil. Some representatives are fungi of genus Fusarium, they cause different plants’ illnesses — fusarium wilt disease of lint, cereals, cotton-plant. 2. Melanconiale. Representative — Gleosporium, cause plants’ disease — anthracosis of pinto, grape, currant and others. 3. Sphaeropsidales. Some representatives are fungi of genus Phoma. Cause rottenness of roots and bulbs of beet and potato, especially dangerous in vegetable store cellar. 4. Mold fungi — Penicillium Aspergillus.

A B Figure 10. Representatives of Deyteromycota: A — Penicillium; B — Aspergillus. Conidiophores and conidium are showed 20 Ecologic groups of Fungi: 1. Soil-inhabiting fungi — mineralize of organic compounds, producing humus — blewits, mycorrhiza-producers; coprophilic fungi grow on the vegetable soil reach of humus. 2. Predatory fungi may be saprophytes and feed on nematodes — small roundwarms. 3. Xylophyles — destroyers of wood, domestic fungi are among them. 4. Water fungi 5. Fungi-parasites of animals and plants. Meaning of Fungi. Fungi play a huge role in a circuit of substances in nature in the reduction of animals and plants, got into soil; creating organic substance in the soil; fungi increase soil fertility; they are widely used in nutrition, for getting the antibiotics, citric acid. Fungi of genus Fusarium produce growth- substance gibberellins, which are used in the agriculture. Fungi are used in medicine. Sclerotium of ergot (Claviceps) have been used for a long time. Therewith, researches of the last years discover a presence of BAS (biologically active substances — vitamins, microelements) in champignons, honey fungi, milk mushrooms and others, which explains their industrial cultivation (champignons). It is also discovered that fly agaric along with the famous alkaloid muscarine contains a variety of highly valued medicinal substances possessing anti- inflammatory and antiseptic activity. Boverin and trichodermin are obtained from the fungi-parasites of insects and are used in insect control. Fungi are widely used in the baking and wine production. However, they also cause harm: parasites of plants cause the loss of harvestand spoiling agriculture products. Fungi cause harm to the forestry, destroy wood conduits. Many of them harm the health of people and animals. Excitans of skin diseases (favus, lichen, dermatitis) are known; fungi may be the reason of chronic maxillary sinusitis. Mycotoxicosis, connected to the toxication by fungi toxins of provisions are very dangerous. The Division of Lichens — Lichenes (Lichenophyta) — Lichen is a com- plexorganism, which is formed as a result of symbiosis of alga and fungus. Fungus gets from alga the organic compounds — carbohydrates, and at the same time fungus provides alga with life environment, defense from overdrying or overheat. Fungus provide alga with the necessary amount of water and water- dissolved mineral salts. Therefore, although the fungus in some ways parasitizes on the alga, but the alga gets a certain benefit from it. Alga provides the fungus with organic nutrition. As a result of these complicated relations a new independent complex organism appeared in the process of evolution. It has characteristic structural features and habitat of life. Vegetative body of lichen thallome is very various by form and colouring. Three types of lichen thallome are separated by external appearance: cristose (or scablike), foliose and fruticose lichens.

21 Thallome of cristose lichens represents crust, strongly fascinated with a substrate — bark, surface of cliffs and stones. This thallome cannot be separated from substrate without damaging it. Thallome of some cristose lichens grows into the substrate, and only fruiting bodies of lichen fungus are seen on the surface, e. g. Xanthoria parietina. Foliose lichens have a form of squama or plates. Their thallome is attached to the substrate with hypha’s batch or rhizoids (Parmelia, Lobaria). Fruticose lichens grow together with substrate only by their basis, their thallome consisting of branches and branched thick stipes (Cladonia, Cetraria islandica). Algae that are part of the thallome are related to The Division of Green Algae and, sometimes, The Division of Cyanobacteria. There are species from the genus of Chlorella, Cistococcus, Cladophora, Pleurococcus) from one-cell green algae in the lichens. The most common representatives of Cyanobacteria are from the genus of the Nostoc. Fungi that are part of the lichens are related, in general, to the Classis of the Sac fungi. Their hyphas, as a rule, form the basis of the lichen’s thallome. Anatomical structure. Thalloms are devided into two types: homeomerous and heteromerous. In a more primitive thallome, homeromerous, alga’s cells or fibers are equispaced between fungi’s hyphas. Slime lichens, containing filamentous Cyanobacteria, have this type of the thallome. Heteromerous thallome has an upper cortex, which is formed by a tight entanglement of fungus’ hyphas. Then a gonidial layer is situated, it consisting of alga’s cells. Fungus’ hyphas, entering the gonidial layer, form small furcations (embranchments), which join the alga’s cells tightly. Here the fungus receives carbohydrates from the photosynthesizing alga. Then lichen’s core is situated — it is a layer of crumbly intertwisted hyphas of the fungi, with their help a certain humidity and aerial environment which are necessary both for alga and for fungus, are supported. After the core, a lower cortex which consists of tightly intertwisted fungus’ hyphas is situated. Rhizoides are marked on the 3a picture. Lichens reproduce in a few ways: 1. Both components reproduce autonomously. 2. By soredia (brood-bad), which represent minute formations, consisting of one or two alga’s cells, surrounded by fungus’ hyphas. They germinate and give the beginning to a new thallome (thallus), in favorable conditions. 3. By isidia, which represent simple or branched outgrowths, fleece an upper side of thallome. They are covered by a cortex from the outside and contain algae and fungi’s hyphas inside. Isidia easily break off from the thallome’s surface and spread with the help of wind and rain, new thalloms are formed in favorable conditions.

Figure 11. Lichens’ structure: 1 — Homeomerous thallus (a — upper cortex, b — algae’s layer, c — the core, d — lower cortex); 2, 3 — homeomerous thallus

Soredia and isidia are characteristic for foliose and fruticose lichens. Many of lichens reproduce by parts of thallome, which are easily broken off by wind or animals in dry weather and spread. This type of reproducing is more typical for cristose lichens. There are known about 25 thousand of lichens. They are widely-spread all around the Earth. The most spread and important lichens are Cetraria islandica (Iceland moss), and Cladonia alpestris (reindeer moss) — the dominant specie in cold desert and forest zone. Foliose lichen from the family Lecanoraceae is very interesting lichen’s representative. It has a shape of round or angulate grey nubbles. This lichen is spread in Asian and African deserts. It is easily come off by wind, lifted in the air and spread for long distances. Sometimes it falls like rain on the ground. It is used for food , That is why it got the name ―manna‖ and became a cause for different legends. The role of lichens. Their meaning in the nature is in being the flora pioneers. Developing on a desert substrate, they prepare it for higher plants. Lichens have a great meaning in feeding reindeer. They are used in getting pure medicinal glucose, nutritional sugar, alcohol, gellatants. In a result of fungus and alga cooperation, specific substances are formed. These substances nowhere else exist. There are so called lichenous acids, some having an antibiotic action. Some lichenous substances act as stimulators, uplifting tonicity of the body. Using decoction of Cetraria islandica in a traditional medicine is based on this property, it is also used as an expectorate remedy. Some of lichens are used in perfume manufacturing, because they have a nice flavor. Conclusion. Fungi and lichens represent a large and underexplored, from the medical point of view, group of organisms. They have interesting biological properties and chemical composition. 23 LECTURE № 3 KINGDOM OF PLANTS (VEGETABILIA OR PLANTAE). HIGHER SPORE PLANTS. COMMON CHARACTERISTIC, CLASSIFICATION, REPRESENTATIVES

The most specialized forms of autotrophic organisms. Zygote gives a beginning to a pluricellular corcle, which develops into a sporophyte (Embryophyta), as their body is divided into stem, leaf and root; they are called Cormophyta. They have a typical cell structure, reproductive process occurs by oogamy. They unite more than 300 thousand species, some of them in a fossil form. They include 9 divisions: 1. Rhiniophyta. 2. Zosterophyllophyta. 3. Bryophyta or mosses. 4. Lycopodiophyta. 5. Psylotophyta. 6. Equisetophyta. 7. Polypodiophyta or Fern. 8. Pinophyta (Gymnospermae, Gymnosperms). 9. Magnoliophyta (Angiospermae) or flowering plants. Rhiniophyta and Zosterophyllophyta are extinct plants’ divisions. The other divisions include both extinct and exiting species. By a number of species and specific features of reproduction, plants are divided into spore and seed plants. In spore plants processes of sporogenesis and gametogenesis proceed on different organisms and at different times. In seed plants gametophyte is reduced and it developes on sporophyte. First terricole plants were Rhiniophyta. They had relative simple structure. They represented not big, bipalmating stems with leaf-shaped outgrowths (awl- shaped or cone-shaped), which had stomatal mechanisms. Primitive, in shape of simple sacs, sporangia developed on the endings of stem branches. There were rhizoides, instead of roots, bipalmating under thin branches. Rhiniophyta became extinct. Only fossil forms (Cooksonia, Rinia) give us an opportunity to judge about the structure and life of ancient ancestors of terricole plants. Plants’ life on the ground made conditions for multicellularity of their organism, differentiation of their cells into tissues, forming specialized organs — stems, roots, leaves, complication of sexual reproduction’s organs, providing developing of corcule, brightly expressed alternation of generations of core phases and interchange of sexual (gametophyte) and asexual (sporophyte) generations. In addition, in most plants (except Bryophyta) sporophyte dominates. In most of representatives, reproductive process depends on a water environment.

24 Higher plants are represented by a large variety of forms. They count more than 300 thousand of species and occupy a dominant position on the Earth. Higher archegonial and flowering plants are separated. Spore and seed plants are found among archegonial plants. First in an evolutional rank stand higher archegonial spore plants. Forming an ovum in archegonium and forming one-celled spores is characteristic for them. The Division of Bryophyta. Represents one of the earliest primitive links at the complicated and long chain of higher plants evolution. Gametophyte prevails over sporophyte. They have a wide spreading — from the Arctic Region to subtropical zone, go up high the hill. Bryophyta are hardly adapted for living in dry places. They inhabit conditions of high humidity — forests, moors, damp meadows, and etc. Sizes of plants are not big, under 20–30 cm. They have chloroplasts and are capable of photosynthesizing. They are autotrophs. The Division of Bryophyta is divided into 3 classes: Hornworts (Anthocerotopsida), Liverworts or Hepatics (Marchantiopsida, Hepaticopsida) and Bryopsida. The Class of Hepatics — Hepaticopsida. Their body is represented by thallus in a form of lamella with a dorsoventral structure. The underside (ventral side) is covered by many one-cell rhizoids, attaching plant with a substrate. A representative of theirs is Marchantia polymorpha (or common liverwort, or umbrella liverwort). Its thallus is covered with epidermis. Lower side’s cells form outgrowths which are called rhizoids. Upper epidermis has breathing openings which is called stigma. Assimilative tissue with large air-filled cavities joins to the upper epidermis; storage tissue joins to the lower epidermis. The plant is heterosexual. Female stalks with archegonia are developed on female species; male stalks with antheridia are developed on male species. Anciently, Marchantia polymorpha was used for treating liver diseases.

A B Figure 12. Marchantia polymorpha: A — male thallus with stalks; B — female thallus with stalks and proliferous baskets 25 The Class of Bryopsida (Musci). These plants have well-defined stems and leaves. Rhizoids are pluricellular and strongly branched. The class includes 3 subclasses: Sphagnidae, Andreaeidae, Bryiidae. The subclass of Bryiidae (The green mosses). They are generally spread in places with excessive or adequate humidity. The typical representative of Bryiidae is moss which is called Polytrichum commune (Common haircap moss). It is widely spread in pine woods, meadows, moorlands; forms bushy turfs. Its stem is under 20 cm in height, covered with hard leaves, cuneal- lanceolar-shaped. Rhizoids begin from an underground part of the stem — rootstock (rhizome). In a central part of stem a primitive conducting bundle is situated. From outside, the stem is covered with epidermis. Life cycle. Plant is dioecious (digamous). On the top of stems of the female species archegonia are formed, antheridia are formed on the top of male species’ stems.

Figure 13. Life cycle of Polytrichum commune

Mosses’ female reproductive organ archegonium is bulb-shaped. In a lower wide part of it — in belly — female gametal cell — ovum is formed and over it a little coeliac canalicular cell. Upper narrow part — collet has a channel with 26 one range of little canalicular cells. Then, canalicular cells fade away into mucus and provide transportation of sperm cell to ovum. Mosses’ male reproductive organ — antheridium has a shape of a long sac with a one-layer wall. Small spermatogenous cells are situated inside the sac. Each of them gives one or two spirally arched sperm cells with two long ciliae. Tops of antheridia open, sperm cells go out from it and they can reach archegonia in a dew or water drop. Then, they go through the collet’s channel to ovum, fertilizing it. Fertilized ovum covers with a shell, divides and capsule on podium is formed. It is called — sporogonium. Its cells have diploid chromosome set. Inside the capsule-sporogonium archesporium is separated. Its cells divide numerous times and after reducing division (meiosis) small haploid spores, serving for asexual reproduction, are formed. Capsules on the tops open, spores are poured out, spread by the wind and germinate in favorable conditions, giving a thin fibre which is called — protonema. Buds are developed on the protonema, each of them may give a moss. Moss also may reproduce vegetative by a rootstock, parts of thallus and etc. In such a manner, spores, protonema’s cells, moss itself, its reproductive organs (archegonia and antheridia) have cells with haploid chromosome set and represent a sexual generation — gametophyte. Diploid phase starts from zygote and ends with cells’ division of archesporium. Therefore, sporogonium (capsule on podium) is sporophyte, an asexual generation of moss. However sporophyte is divided, but not divided from gametophyte and feeds at its expense, though it can photosynthesize by itself. Sporophyte dies out the same summer, after spreading spores. Other Bryiidae mosses — Ptilium, Dicranum, Mnium, Holocomium form continuous cover of mosses in the northern pleurocarpous moss forests. The subclass of Spagnidae unites over 300 species that are spread in the northern forest zone on big swamped spaces. Spagnidae have a stem and leaves like phyllodes. Stem and leaves have dead cells with ringed and spiral thickening of membranes. Small living chlorophyll-containing cells are situated between them in the shape of net. Dead cells are filled with air, which forms on them whitish colour and determine their huge hydroscopic properties. One part of dry moss absorbs 25–30 parts of water. The representative is Sphagnum magellanicum. Spagnidae form peat-bogs. Peat is widely used as a fuel material, fertilizer, and stock material in a chemical industry (alcohol), sealer and insulant. It is used in medicine in the quality of hydroscopic material for dressings, pads, separators, and etc. The Divisions of Lycopodiophyta, Equisetophyta, Polypodiophyta. They represent a large group of higher land spore plants, having the leaves, stems, and roots with fibrovascular bundles. In their life cycle the alternation of

27 generations is well-defined. Sporophyte is represented by a vegetative plant. Haploid spores are formed on sporophyte in sporangia. In some representatives — homosporous — they are morphologically equal, in other representatives — heterosporous — spores are morphologically different — microspores and macrospores. Prothallus (sexual generation) is developed from spores. In homosporous it is unisexual, in heterosporous it is heretoecious (male and female).

Figure 14. Sphagnum sp.: A — general view; B — top of stem with sporangium; C — moss-capsule; D, E — phyllode in a lateral cut; F — stem in a lateral cut. 1 — false podium of moss-capsule; 2 — moss- capsule; 3 — podium; 4 — column; 5 — sporogenous tissue; 6 — wall of sporangium; 7 — chlorophyll-containing cells; 8 — dead water-containing (hyaline) cells; 9 — conductive cell; 10 — mechanical cell; 11 – ―bark‖

The Division of Lycopodiophyta. Tree-like (arboraceous) forms are extinct; they had an important meaning in formation of the fossil coal. Herblike plants of small size (low size) — fewer than 70 cm with low-branched roots and not big thin leaves like phyllodes are remained. Both homosporous and heterosporous forms are among them. Two classes are separated: Lycopodiopsida and Isoѐtopsida. The Class of Lycopodiopsida. Homosporous forms are represented by plants of the Lycopodiales order. Their typical representative is Lycopodium clavatum, which is spread in soft-woods forests (pine woods). It has climb, dichotomically branched stem with small line-awl shaped leaves and low-

28 branched roots. Stem’s length is under 6 and more meters. There are sporogenous spikelets gathered in pairs on apex shoots. Spikelet consists of stem with leaves — sporophylls (leaves which bear sporangia). Single bud-shaped sporangia with a big amount of equal spores are situated in sinuses of sporophylls. Spore produces prothallus — a bisexual lycopodium’s gametophyte. Prothallus is chlorophyll-free, braided with fungi’s hyphas — mycorhiza. On the upper side of gametophyte, archegonia with ova and antheridia with sperm-cells are developed. Corcule is developed from zygote after fertilization; adult plant of sporophytic type is developed from corcule. Formation of gametophyte from spore and developing sporophyte from it takes more than 20 years. Lycopodium’s spores are used in medicine as a baby powder, for filling pills; in foundry engineering and etc.

Figure 15. Lycopodium clavatum: A — sporophyte; B — sporophyll with sporangium; C — spore

The Class of Isoёtopsida. Heterosporous form is represented by Selaginella selaginoides of the class Isoѐtopsida, which is spread mostly in the tropics. In the Commonwealth of Independent States (The CIS), only 8 species are vegetated. Selaginella has more tender dichotomically branched stems of a small size. Forms small shrubs fewer than 10 cm. Sporogenous spikelets have sporangia of two types. There are microsporangia with a big amount of microspores and megasporangia with 4 megaspores. Heterospory, as a beginning of sexual differentiation, represents a progress in evolution. Spores produce prothalli: microspores which are male, and megaspores which are 29 female. Gametophytes are reduced very much. Male prothallus consists of antheridium and one vegetative cell; female consists of a few archegonia with ova. Prothalli are very small and they don’t leave spore’s cover. Corcule is formed from zygote after fertilization; an adult sporophyte is formed from corcule. These plants do not have big practical meaning. Sometimes tropical forms are cultivated as decorative plants.

Figure 16. Selaginella selaginoides: A — sporophyte: 1 — an additional root, 2 — stem with leaves, 3 — strobilus, 4 — young sporophyte; B — strobilus with sporangia, 5 — microsporangium, 6 — megasporangium, 7 — microspore, 8 — megaspore

The Division of Equisetophyta. Tree-like representatives of this division — calamites — played a big role in the vegetation cover of the Earth in the Carbonic period. Only small herblike plants of the genus Equisetum have remained, accounting about 20 species. Equisetum arvense, equisetum pratense, equisetum palustre, equisetum sylvaticum, equisetum ramosissimum are spread in the CIS. We will explore specifics of biology and plant’s structure on the example of Equisetum arvense. As a rule, it is perennial field weed. It winters it exists in a shape of rootstock with a store of nutrient materials both in a rootstock and in tubers, which is formed on rootstocks. Whity-pink sporogenous shoots with fewer than 20 cm high grow from rootstocks in early spring. Stems of these shoots are erect, juicy, not branched with strongly-expressed nodes and internodes. Leaves are underdeveloped, small, and narrow; they grow together in a tube by their bases. Leaves are situated by whorls. Sporogenous spikelet is on shoot apex. Axis of sporogenous

30 spikelet is covered with changed leaves — sporogenous leaves. Sporangia are formed on sporogenous leaves, spores are formed in sporangia. Outwardly spores are equal, but physiologically they may be different. Each spore is crossed by 4 ribbon-like appendages — elaters, keeping a few spores together. After maturation of spores, ribbons are straightened and overspread spores.

Figure 17. Life cycle of Equisetum arvense: A — vegetative shoot; B — sporogenous shoot; C, D — leaf sheath; E – sporangiophore; F–H — spores: 1 — rootstock with tubers, 2 — strobilus, 3 — corymb (fascicle), 4 — sporangium, 5 — podium, 6 — an external spore’s cover, 7 — elaters

In favorable conditions spores will germinate and will give bisexual or dioecious prothallus, which depends on conditions of germinating. Archegonia and antheridia are submerged into corcule’s tissue. About 200 sperm cells are formed in antheridia, which can move in a water drop. Zygote and corcule are developed from fertilized ovum; an adult sporophyte is developed from corcule. After maturating and falling of spores, spring sporogenous shoots of horsetails die away. Instead of them, vegetative shoots of sporophyte are formed from the same rootstock. Stems of these shoots are erect, branched with reduced squamose leaves. They provide refilling of storage materials in rootstock. In such a manner, sexual and asexual generations exist independently. The horsetails (Equisetum) do not have big practical meaning. Vegetative shoots of Equisetum arvense are used in a medicine as diuretic agent. The Division of Polypodiophyta (Fern, Pterophyta) Ferns, as well as lycopodium and horsetails (equisetum) derive from Psylotophyta. The division

31 unites about 10 thousand of species. There are 100 species of the division in the CIS. It includes classes ofMarattiopsida, Polypodiopsida and others. In the quality of a typical representative of Polypodiopsida we will explore Aspidium filix mas (male fern), which is widely spread in the forests of temperate zone. It reproduces by rootstocks, by sexual and asexual ways. A fascicle (a bunch) of leaves grow from rootstocks. Young leaves are hook- shaped, coiled in the bases of adult ones, they start to uncoil under the soil and complete the formation during their third year, achieving one and more meters in length. The formed leaves are bipinnatiparted, grow by their tip and die each year.

Figure 18. Life cycle of fern

Sporangia are formed by the midsummer on the lower side of a leaf. Sporangia are placed in groups on a thick outgrowth — placenta, forming sori. Spores of a round shape with a double cover (exine and intine) are situated in each sporangium. In favorable conditions spore germinates and gives a prothallus (sexual generation, gametophyte). On a lower side of the prothallus, rhizoids are developed, by which the plant is attached to the ground; higher there are antheridia and archegonia. Moving of sperm cells is conducted through water. 32

Figure 19. Salvinia natans (floating fern): A — general view of the plant from above; B — micro- and megasori (in section)

A corcule grows from zygote; an adult plant — sporophyte (diploid) — grows from the corcule. Prothallus perishes. Except for male fern, common bracken (Pteridium angelinum) and female fern (Athyrium filix-femina) grow. From the point of view of the evolution’s progress, a water fern — Salvinia natans is of some interest, the fern produces spores of two types: microspores and megaspores in micro- and megasporangia. It is the beginning of sexual differentiation. Ferns do not have any big practical meaning. Rootstocks of Aspidium filix mas (male fern) and of some other species are used in a medicine for preparation of antihelminthick preparations. Conclusion. Two directions are separated out in evolution of a higher spore plants. In the 1st direction, gametophyte is developed and improved, and sporophyte takes a secondary place. It is a line of mosses (Bryophyta), which did not result in the creation of new plant forms. It turned out to be a blind line in evolution. In the other line, sporophyte is complicated and takes a dominant place in a cycle of plants’ developing; gametophyte is reduced all the more. The evolution of a sporophytic line went through the developing of macrophilia, heterospory, dividing body into organs, improving and complicating of stem, gradual rejection from fertilization in water environment. It is the reason of all the variety of vegetal life on the planet.

33 LECTURE № 4 DIVISION OF GYMNOSPERMS. GENERAL CHARACTERICTIC, CLASSIFICATION, REPRESENTATIVES

Gymnosperms (Pinophyta/Gymnospermae) are higher archegonical seed plants. Seed ferns are thought to have been stirs of Gymnosperms, which died out at the end of Carboniferous period because of cooling. Unlike cryptogamous plants, Gymnosperms multiply by seeds, which develop in ovule. Seeds have embryo and endosperm (stock of nutrients for embryo that develops from zygote). Endosperm is formed from megaspore before fertilization (maternal origin). It plays a big role in plants’ survival. Ovule’s covers (that are called integuments) protect an embryo from unfavorable conditions and then become seed coats. Seeds do not have a pericarp and are located on cone’s scales. The presence of embryo, endosperm and integuments are the most important adaptations of Gymnosperms to terrestrial lifestyle. Gymnosperms’ structure. Usually they are represented by arborescent or shrubby forms. They have a main root with tubercles or mycorhiza, leaves (fir- needles/needles) are minute, numerous (unlike the ferns). Only sago-falms preserve large leaves. Needles live for 2–5 years, and only a part of them shanks. That is why plants are evergreen. A stem has monopodial branching. There are 2 types of sprouts: 1) lengthened (with squamules) 2) shortened (are formed in axils of squamules) — as a rule, they are covered with leaves. Sprouts end up with cones or strobili. Strobili, as a rule, are unisexual, and plants are monoecious(sago-falm is dioecious). Male strobili consist of an axis and microsporophylls that are formed at the base of young sprouts, in axils of squameles (instead of shortened sprouts). Female strobili are formed at the top of lengthened spring sprouts. Fertilization of the most ancient representatives (sago-falms) is implemented by spermatozoids that move with flagellums in liquid pollen tube; at present-day plants — by sperms without flagellums that move to ovum through pollen tubes. So their sexual process is independent from aquatic environment. Gymnosperms have an observed generation change: sexless generation is the whole plant; sexual one is much reduced male and female gametophytes. They develop apart: female gametophyte is formed from megaspore inside the ovule, which is the modified megasporangium; male gametophyte is sprouted pollen, an antherid is totally absent. Gymnosperms are wide-spread around the Globe. Classification is still unstabilized. More often 6 classes are divided: 1) Seed ferns 2) Cycads

34 3) Bennettitales 4) Gnetalians 5) Ginkgoales 6) Conifers Seed ferns (Pteridospermae). This is an extinct class. They are important only as fossils. Seed ferns came about from heterosporous ferns. They are a transitory team from ferns to gymnosperms. Structure and leaves’ outlook are similar to ferns and presence of seeds and ovules, secondary growth are common with gymnosperms. Class Cycads (Cycadopsida) unites about 20 species of present-day plants. Cycads are distributed in tropical and subtropical regions of Eastern Asia, Australia, Africa and America. They are slow growing trees up to 20 m height living up to one thousand years. Stem is not branching, it is column-like. Cycads have large pinnately leaves 3 m long. The core is very developed, where large amount of starch is stored. Since the olden pre-historicage the plants are used for production of groats. They have large strobili; female ones (about 10 kg) are used as food; male strobili are united in small groups. Fertilization is not associated with water. Spermatozoids have many flagellums. The lants are wonderfully decorative. One of the Representatives is Cycas revoluta.

Figure 20. Cycads: 1 — Cycas revoluta; 2 — Stangeria paradoxa; 3 — Zamia floridana

Class Bennettitales (Bennettitopsida). Plants are totally extinct. They had bisexual cones. Structurally they were similar to Cycads. Class Ginkgoales (Ginkgoоpsida) is represented by a single species — Maidenhair-tree bilobular(Ginkgo biloba). This is an evergreen tree up to 40 m heigh with branching dense pyramidal crown. Leaves are stalked with a fan- shaped blade. It’s a dioecious plant. Microsporophylls are located spirally on the catkin’s axis. Microspore-cases are located two by two at both ends of each 35 catkin. Megasporophyll is tubercle-shaped and has 2 ovules. Pollination occursby wind. Spermatozoidis are polyflagellumous.

Figure 21. Maidenhair-tree bilobular: A — shortened sprout with leaves and microstrobilum; B — shortened sprout with leaves and megastrobilum; C — seed; D — leaf

The Seeds are large, looking like stone-fruit and are used as food. Its Wood is valued as ancient conifers. It’s habitat is South-West China. The plant is widely cultivated. It is known as a sacred tree in China and Japan. The plant is represented as a decorative on in many botanical gardens. The leaves are used for preparation of some medications. Class Conifers (Pinopsida) is Nowadays the most numerous representative of Gymnosperms. There are 600 species. Class is mostly spread in the Northern hemisphere, where extensive conifer forests are formed (pine, spruce, larch, fir). In general variety of conifers is concentrated around the perimeter of the Pacific Ocean. Conifers appeared long before formation of modern mountains and plains. It is considered the China’s center of origin to be the ancient one. Modern Conifers achieved high xeromorphic structure unlike the other Gymnosperms, which is why they are so successful. In winter despite the presence of needles, conifers evaporate much less water than Angiosperms. It is a progress of evolution in microphyllous line. Grassy forms are not known. There are only trees and shrubberies. Branching is monopodial, rarely sympodial. Conifers are distinguished by existence of needles, absence of spermatozoids. The majority of representatives are monoecious, instead of dioiceous. Male cones are much smaller than female ones. Seeds are provided by scarious winged caruncle.

36 Subclass Conifers includes 7 orders. The most important are Pines, Yews, Cypressed. Order Yews (Taxales). Yew (Taxus orientale) is a wide sprawling, shade- loving, beautiful tree with several tips. The plant makes a high-quality wood. Now it is a rare plant. It preserves on the Caucasus, Carpathians, in the Crimea and in Belarus. Order Pines (Pinales). The most extensive order that has large area and is represented in all continents. Leaves are alternative, single or in bunch. The plants are monoecious, sometimes dioecious. Cones are woody, dehiscent at ripening. Pollen has air sacs (rarely without). Order includes 10 families, inhabiting mainly in North hemisphere (pine, spruce, larch, fir, cedar and so on). Scotch pine (Pinus sylvestris) is a straight-stemmed tree up to 50 m heigh, having by 2 needles in bunch and themain root. Cones are solitary or located by 2–4 near sprouts’ tips. It’s a light-loving plant. There are schizogenous resin ducts, containing essential oils, resins, balsams, in bark, wood and leaves.

Figure 22. Developing cycle of conifers (pine): 1 — male cone; 2 — microsporophyll with microspore-case; 3 — pollen; 4 — female cone; 5 — megasporophyll; 6 — squamule with 2 seedbuds; 7 — scale with 2 seeds in a triennial cone; 8 — germ

Pine is a monoecious plant. Microsporophylls are united in cones, sitting at a base of young branches. A cone has an axis. So, microsporophylls circumferentially sit on it from top to bottom. Each microsporophyll has 2 microsporangiums on the lower side, where microspores develop. Microspore still germinates in microsporangium. During division of its nucleus one antheridial cell and one vegetative one are formed. Vegetative cell

37 dies out and antheridial one forms a basal cell and generative (spermatogenous) cell, which gives 2 sperms. After formation these 2 cells microsporangium opens and pollen are spread by wind. Female cones appear at the top of young sprouts. 2 types of scales are located on the cone’s axis: outer sterile, which are called protective scales, and inner seminal. On the top of seminal scales 2 ovules (megasporangies) are located. Protective scale is a sterilized megasporophyll. Ovule consists of nucellus and integuments. Micropyle is situated on the top of ovule. Usually one of the nucellus’ cells becomes maternal cell of megaspores. It divides with reduction of chromosomes into 4 megaspores. 3 upper ones of them degenerate and the last one preserves. Growing up this megaspore gives female multicellular prothallium that is called endosperm. On the prothallium’s side, which is faced to the micropyle, 2 archegonia are formed. In each archegonium one large ovum and 2–4 small neck cells are polarized. Landing through micropyle on the ovule’s nucellus, pollen for a long time (1 year) sleeps (rests), clinging to it. In 1 year pollen germinates. Basal cell forms pollen tube that intrudes in nucellus and grows in its tissue. Antheridial cell divides into two: generative and vegetative cells. The last one comes to the tube and provides the environment for the generative cell’s promotion. Generative cell gives 2 sperms (not spermatozoids). Pollen tube transfers them to archegonia and bursts. One of sperms permeates to ovum and fertilizes it. Zygote divides, giving an embryo that develops at the expense of primary endosperm (that has maternal origin). Seeds’ ripening occurs at the end of the second year. Female cone grows, changes its color from red to green. During seeds’ developing and fertilization scales link one to each other. Mature cone becomes brown, scales open and seeds spill out. So, mobile spermatozoids are not formed, sperms are instead of them. Sperms are transfered to archegonia by pollen tube. Finally, sexual process is independent from water environment. Pine’s buds as an expectorant remedy and cones of the common spruce (Picea abies) as an antiphlogistic one are mentioned as a medicinal raw material in the State Pharmacopeia of the Republic of Belarus. Order Cypresses (Cupressales) is represented by trees and shrubberies. Leaves are opposite or whorled, scaly or needle-shaped. Genus Juniper (Juniperus sp.) The representative is common juniper (Juniperus сommunis) (red, sibericum, savin and etc.). Fleshy female cone’s scales grow together, forming cypress cones, which are included as a medicinal raw material in the State Pharmacopeia of the Republic of Belarus and called ―Juniper’s fruits‖. They are recommended as a diuretic remedy. Genus Cypress: Cypress evergreen (for green building).

Figure 23. Juniperus сommunis

Class Gnetalians (Gnetopsida). It is represented by monoecious or dioecious plants which have a single ovule with 1–2 envelopes, elongated in a tube at the ends. A tube looks like a style of pistillate plants. Secondary wood has vasculars, there are no scalariform tracheids and resin ducts; leaves are opposite, embryos are bilobate. These features are characteristic of this class. It includes 3 orders: Ephedras, Welwitschiales, Gnetales. 1. Ephedras (Ephedrales). Ephedra equisetina, or ephedra horse-tail. It is a small shrubbery belong 40 cm heigh, looks like a horse-tail and is spread in Europe and Asia. Green ribbed articulate stem carries inconspicuous scaly opposite leaves. The plant is dioecious. Strobili look like inflorescence ―spikelets‖. Ovule is enclosed in a special cover, which grows after fertilization and reminds juicy pericarp (like stone-fruit). It contains ephedrine (2–3 %). The plant is used for cardiovascular diseases, nervous system diseases, bronchial asthma and etc.

Figure 24. Ephedra horse-tail, the branch of a flowering pistillate plant: а — stamen’s spikelet (congestion of microstrobila); b — a seedbud with sterile protective scales; c — seed 39 2. Welwitschiales (Welwitschiales). The order is represented by 1 species — Welwitschia mirabilis — Welwitschia miracle that preserves as an endemic plant in Angola, South-West Africa. It looks like a giant germ with two opposite leaves 2–3 m wide. Conifers’ value. Cone-bearing plants are the most valuable. Pine or spruce wood is used as building material, paper; larch bark contains 10 % of tannins; cedar nuts — 50 % of fixed oil; pine needles have vitamin C; essential oils, resins and balsams are used to produce turpentine, rosin, wood vinegar; drug- purpose. Evolution value of Gymnosperms. Gymnosperms are a big step in evolution of Flora. A range of features, which lets Gymnosperms take valuable place in Flora, appear to be the presence of ovule, seeds, differentiation on organs; conductive system represented by vasculars; domination of a sporophyte (reduction of gametophyte); sexual process not associated with water.

40 LECTURE № 5 ANGIOSPERMS. GENERAL BIOLOGICAL CHARACTERISTIC. MORPHOLOGICAL DIAGNOSTIC FEATURES OF VEGETATIVE ORGANS

Angiosperms (Magnoliophyta / Angiospermae) are higher flowering plants. They occupy a dominant position in vegetation cover of the Globe. They appeared in Cretaceous period of Mesozoic Era. There are about 250 thousand- 300 thousand species nowadays. Unlike previous divisions of Flora: 1. They are represented by trees, bushes, herbs (annual, biennial or perennial one). 2. Vegetative organs consist of very various histological elements. Also Angiosperms have vessels. 3. Angiosperms have specialized vegetative organs, such as root, stem, leaf, rhizomes, tubers, bulbs. 4. They have true flowers. 5. There are pistil and fruit (seedbuds are located under carpels’ cover), seeds are covered by pericarp. 6. Further reduction of male and female gametophyte takes place. 7. There is a Double fertilization. 8. Endosperm is hybrid, triploid. 9. Embryo has 1–2 seed-lobes. Angiosperms are on the top of complex chemical composition and variety of flowing physiological processes. They play a big role in human economic activity acting as food, clothes, habitation, fuel, remedy. The entire evolution of modern Kingdom of Animals, especially mammals, birds and human’s appearance, is associated with flowering plants.

THE REQUIREMENTS OF THE STATE PHARMACOPEIA OF THE REPUBLIC OF BELARUS TO THE EXTERNAL STRUCTURE OF VEGETATIVE ORGANS OF PLANTS Leaves (Folia) are medicinal raw material in form of dried out or fresh leaves or single leaflet of a compound leaf. Macroscopic diagnostic features: – The type of leaves (simple or compound). – The form of blade and leaf stalk (fig. 25). – The feature of leaf edge (fig. 26). – The insertion way of a leaf to stem (fig. 27). – The type of dissection of blade (fig. 28). – Venation (fig. 29). – Pubescence.

Figure 25. The form of blade: 1–3 — rounded; 4 — oval; 5 — oblong; 6 — lanceolate; 7 — oblong-ovate; 8 — inversely lanceolate; 9 — elliptical; 10 — rhomb-shaped; 11 — ovate; 12 — ovate-heart-shaped; 13 — broad ovate; 14 — trigonal; 15 — heart-shaped; 16 — inversely ovate; 17 — oblong inversely ovate; 18 — linear; 19 — spathulate; 20 — nodular or kidney-shaped; 21 — arrow- shaped; 22 — spear-like; 23 — needle-shaped; 24 — fan-shaped; 25 — with oblique base

Figure 26. The form of a tip and base, the feature of leaf edge: А — tip: 1 — rounded, 2 — flat; 3 — acuminate; 4 — sharp; 5 — sharp-pointed; 6 — obtuse; 7 — emarginated; 8 — bilobular; B — base: 9 — rounded; 10 — cuneated, 11 — flat; 12 — heart-shaped; 13 — arrow-shaped; 14 – spear-like; 15 — oblique; 16 — narrowed; C — edge: 17 — entire; 18 — toothed; 19 — serrated; 20 — heteromerous-biserrated; 21 — cremated; 22 — emarginated; 23 — macro emarginated; 24 — ciliate; 25 — wavy 42

Figure 27. The insertion way of a leaf to stem: 1 — leaf blade and it’s parts: a — base, b — tip, c — edge; 2 — leaf stalk; 3 — stipules; 4 — leaf base; A — total leaf; B — short-pedicellate leaf without stipules; C — sessile leaf, attached with base; D — a leaf with pulvinus; E — decurrent leaf; F — amplexicaul leaves; G — perfoliated leaf; H — opposite, connated, perfoliated leaves; I — leaves with sneath; J — leaf with ochrea

Figure 28. The type of dissection of blade: 1–5 — pinnate: 1 — pinnately lobed, 2 — pinnately parted or runcinated; 3 — pinnately sected or lyrate, 4 — unequally — intermittently — pinnately sected; 5 — repeatedly — pinnately sected; 6–9 — palmate: 6 — palmatilobated, 7 — palmatipartite, 8 — palmatisected, 9 — thrice — palmate — pinnately sected; 10 — ternately lobated; a — lobes; b — parts; c — segments

Figure 29. The types of leaf’s venation: 1 — pinnately marginal; 2 — pinnately loop-shaped; 3 — pinnately netted; 4 — palmati- marginal; 5 — palmatiloop-shaped; 6 — palmatinetted; 7 — parallel; 8 — curved

Figure 30. Types of a phyllotaxy: 1 — alternative or spiral (1 leaf in each node); 2, 3 — opposite and crosswise opposite (2 leaves in each node); 4 — whorled (more, than 2 leaves in each node); 5 — rosetted

Figure 31. Leaves’ metamorphosis: 1 — bud and protective scales; 2–5 — stages of modification of leaf parts in tendrils; 6 — tendrils(modifications of rachis and superior leaflets of a compound leaf); 7 — spicules (modified leaves); 8–10 — stages of transformation of leaf parts in spicules;11 — stipules, modified in spicules; 12 — phylloids (flat photosynthesis leaf stalks); 13 — trapping apparatus of insectivorous plant

Herbs (Herbae) are medicinal raw material in the form of dried out or fresh overhead parts of herbaceous plants (stems with leaves and flowers, in a measure with flower buds and fruits). Macroscopic diagnostic features of stems: – The location of a stem in a space – The type of ramification – The phyllotaxy – The form of cross-section – Size (length and diameter at the base) – The features of surface, pubescence.

Figure 32. The location of a stem in a space: 1 — upright; 2 — clinging; 3 — twisting; 4 — climbing with suckers; 5 — creeping; 6 — trailing

Figure 33. The type of ramification: 1–2 — tip branching, forked increase(equal- and anisoforked); 3–5 — lateral branching, monopodial increase with alternative (3), opposite (4) and whorled (5) location of lateral axes; 6–8 — lateral branching, monochasial (6), dichasial or falseforked (7), pleiochasial (8) increase; 9 — tillering; 10 — columnar stem

Figure 34. The form of cross-section: 1 — cylindrical; 2 — elliptical; 3 — rounded tetrahedral; 4 — concave tetrahedral; 5 — trihedral; 6 — polyhedral; 7 — articulate; 8 — grooved 46 Roots (Radices), rhizomes (Rhizomata), tubers (Tubera), bulbs (Bulbi), corms(Bulbotubera) are dried out or fresh underground organs of perennial plants, which are clean from the soil and free from traces of stems and leaves. Macroscopic diagnostic features: 1. Size and form. 2. The features of surface and fissure(fracture). 3. The color of surface on a fresh fissure(fracture). 4. Smell and taste.

THE STRUCTURE OF THE FLOWER. INFLORESCENCES The main distinctive feature of Angiosperms is the existence of a flower. There are some theories of the flower genesis: 1. The flower is a modified sporogenous sprout. The flower is a shortened and limited in growth sprout, where megasporophylls are transformed into carpels and microsporophylls are transformed into stamens; the flower is created to form spores and gametes for sexual process. After it seed and fruit are formed. 2. The flower is a modified inflorescence and all of its parts are accrete single flowers.

Figure 35. The structure of a flower: 1 — stigma; 2 — ovary; 3 — style; 4 — seedbud; 5 — filament; 6 — connective; 7 — anther; 8 — a cutaway anther; 9 — pollen grains; 10 — petal; 11 — calyx lobe; 12 — receptacle; 13 — pedicel; 14 — bract; 15 — bractlet

47 Parts of a flower: – pedicel is a stemmed part of a flower; – receptacle is a wide part of a pedicel. All the other parts are attached to the receptacle. It can be flat, convex, concave, and etc., and also disk-shaped, saucer-shaped, conical; – calyx lobes (Calyx) are an external part of a perianth (Calyx). It can be colored or green; segmented or with united segments. Calyx lobe can be awl- shaped, lance-shaped. Double circle of a Calyx lobe forms outer calyx (calycle); – petals (Cоrolla) form corolla and, as a rule, are colored. They can be free or united, have various forms and sizes. Sepals and petals form a double perianth. A perianth is simple if there are only petals or sepals (Perigonium). A perianth serves: 1) to cover sporophylls; 2) to calling in insects; – Stamens (Androceum) and pistils (Gineceum) are situated on a receptacle inward from the perianth. Stamens are microsporophylls that consist of a filament and an anther. Reduced stamens (without anthers) are called staminoids. An anther consists of 2 equal parts, which are divided by a connective. Each part has 2 antherlobes, where pollen ripens. The number of stamens differs from 1 to a few dozens. They can also be free or united, which is why androecium can be monadelphous, diadelphous, polyadelphous. The inner side of a receptacle is occupied by the gynoecium. Gynoecium is a complex of megasporophylls (or carpels) that forms pistil (1 or several). A pistil consists of an ovary (lower extended part), style and stigma. If there is no style, a stigma is sessile. Forms of stigma iare very various: capitated (plum, merry, cherry); bilobated (valerian); trilobite (blue-bell); stellate (in form of a star), etc. Pistil is formed by a concrescence of megasporophylls. Ovules develop on the inner side of them. Ovary can be superior (hypogynous flower), semi-inferior (halfway accreting with receptacle), and inferior (it means that all flower’s parts are situated above the ovary) (epigynous flower).

Figure 36. Position of ovary: 1, 2 — superior; 3 — inferior; 4 — semi-inferior

Types of gynoecium (fig. 36): 1) monocarpous (from 1 carpel); 48 2) apocarpous consists of several carpels. Each carpel forms its own pistil (the most primitive); ovules are parietal (magnolia); 3) coenocarpous (1 pistil is formed from several united carpels; ovules are located in center or at the edges or united magesporophylls): – syncarpous (compound ovary); – paracarpous (single ovary, parietal placentation); – lysicarpous (single ovary, ovules are in center). Carpels can totally unite in one pistil, from ovary to stigma; sometimes only by ovaries. So, styles are free. Sometimes it occurs by ovaries and styles, so, stigmas are free. Actinomorphic flower is a flower, where it’s possible to hold several axes of symmetry (tobacco, flax, cotton). Zygomorphic flower is a flower, where only 1 axis of symmetry is holding (Pod-beaning family, Mint family, violet). Asymmetric flower has no axes of symmetry.

Figure 37. Types of gynoecium (cross-section) and likely direction of its specilization: 1 — monocarpous; 2 — apocarpous; 3–5 — coenocarpous types: 3 — syncarpous, 4 — paracarpous, 5 — lysicarpous

Formula. The flower’s structure can be expressed by formula or diagram. Parts of a flower submit to the rules of a phyllotaxy. Formula is an alphanumeric expression. (*Са5Со5А5G5) Diagram is a graphic expression of the flower’s structure.

Figure 38. Flower’s diagram: 1 — protective leaf; 2 — calyx; 3 — corolla; 4 — stamens; 5 — pistil; 6 — stem; 7 — ovary

Figure 39. Diagrams of flowers: 1 — acyclic flower; 2 — hemicyclic one (carpels form 1 circle); 3–6 — cyclic: 3 — actinomorphic, pentacyclic, pentamerous, 4 — actinomorphic, quadricyclic, pentamerous, 5 — actinomorphic corolla, but a flower has only 1 axis of symmetry, 6 — zygomorphic (here bract’s place is showing by the dotted lines)

Inflorescences are a group of flowers on a common peduncle. Botrious (monopodial) inflorescences. Main axis does not limit in growth. Flowering occurs in upward order. There are simple inflorescences (raceme, small shield, spike, cob, catkin, capitulum, umbell) and compound ones (compound spike, panicle, compound umbel, compound corymb). 50

Figure 40. Types of botrious inflorescences: А — Simple inflorescences: 1 — raceme; 2 — spike; 3 — cob; 4 — umbell; 5 — basket; 6 — capitulum; 7 — small shield; B — Compound inflorescences: 1 — panicle; 2 — compound corymb; 3 — compound spike; 4 — compound umbel

Simpodial (cymous) inflorascences have limited growth of a main axis: simple cyme (coil, bend), dichasial cyme (Pink family), pleiochasium (Spurge family).

Figure 41. Types of simpodial inflorescences: А — Single cyme: а — bend, b — coil, c — double coil; B — Dichasial cyme; C — Pleiochasium

Thyrsuses are branched compound inflorescences, and measure of branching decreases from base to apex. Main axis grows monopodially, and cymoids are lateral inflorescences (horse chestnut, mullein, inflorescences of Mint family, Composite family, and birch). 51 Pollination is a transfer of pollen to the stigma of a pistil. Self-pollination is observed in hermaphrodite flowers, when pollen spills out on the stigma of its own flower. It is vitally important that pollen and embryo sac must ripen at the same time. The absence of physical or biochemical obstacles is necessary too (for example, the anther opens at the side, which is opposite the stigma). Self-pollinating adaptations: – self-pollination in a close flower bud, cleistogamous flowers (peanut); – grasses have an inflorescence in a leaf sneath; – pollen and ovary(embryo sac) ripen at the same time. It should be mentioned that pollen of self-pollination plants can pollinate another flower of the same species. Xenogamy (cross-pollination) is a pollen transfer from stamens to stigma of pistils of different flowers of the same species. According to the method of pollen transfer plants can be divided into: – anemophilous (birch, alder, oak, hazelnut). Fine nondescript flowers without aroma and nectary. Wind-pollinated plants have a lot of flowers and pollen (fine, dry); – entomophilous. It is a result of phylogenetic development of organisms in conditions of natural selection. Nectar is insects’ food. Nectaries are located in form of grooves, tubercles, spurs , etc. A large amount of pollen is produced. So, flowers are large and bright, single or in inflorescences. Doubleness is a conversion of stamens or pistils into petals. There are no seeds (dahlia, chrysanthemum, double roses). Conclusion. Flower is an organ of a plants’ seed multiplication, which is adapted to provide sexual and sexless multiplication. At macroscopic diagnostic of flowers and inflorescences as medicinal raw material (according to the SP of the RB) type of inflorescence, pubescence, structure of perianth (calyx-shaped, corolla-shaped or double, actinomorphic or zygomorphic), amount and form of calyx lobes (or minute teeth of calyx), the amount and form of petals (or minute teeth of corolla), amount and structure of stamens and pistils, features of ovary’s structure are marking. Sizes of flowers and inflorescences are measuring. Microscopy: preparations from distant parts of inflorescences (flowers, leaflets, spathes) or parts of flower (petals, calyx lobes) are prepared. Viewing their surface special attention is paid to the structure of epidermis, presence and structure of hairs, glands, crystal inclusions, mechanical elements (in spathe’s leaflets), form and size of pollen grains, etc.

52 LECTURE № 6 BIOLOGY OF FERTILIZATION. STRUCTURE AND DEVELOPMENT OF FRUITS AND SEEDS

Microsporogenesis or formation of male gametophyte. An adult stamen consists of anther, connective and filament, which is covered by epidermis. Stamen is formed from tubercle of growing point of floriferous sprout, which is initially homogenous and then differentiates. Gradually in 2 sites of each pollen sac inception of anther nests begins; archeosporial cell originates, divides into parietal and sporogenous cells.

A B Figure 42. Structure of a stamen (А) and anther (cross-section — B): 1 — filament; 2 — anther; 3 — connective; 4 — underconnective; 5 — vascular bundle; 6 — anther nest (according to 1 microsporangium); 7 — dehiscent half of an anther with spilling pollen grains; 8 — anther’s wall

Parietal cells form subepidermal fibrous layer and tapetum that provides sporogenous tissue with nutrients. Cytoplasm is viscous, cells are large. Sporogenous cells divide by meiosis and give microsporecells. In future these cells will differentiate into pollen. The nucleus of microspore divides into generative nucleus and vegetative one. Vegetative nucleus is large, contains greasy oils and starch. Generative one is smaller and has a chromatin. It can divide even in an anther, giving 2 sperms (Composite family, grasses, Madder family, Pink family), or in a pollen tube (Orchid family, Figwort family). Consequently, mature pollen grains can be binucleate and trinucleate; they are covered with sporoderma, consisting of exina and intina. Exina is a thickened tunicate coat with excrescences in form of tubercles, small prickles, etc. Exina consists of special substance pollenin (cellulose impregnated with 53 cutin substance). Exina has pores. The form and surface are its diagnostic features. Intina consists of pectin substances. Opening of anthers occurs during desiccation due to diminution of epidermis cells.

Figure 43. Structure of a pollen grain (scheme)

Figure 44. Forms of pollen grains: 1 — monofurrow (magnolia); 2 — trifurrow (peony); 3 — polifurrow (Hymenocrater); 4 — monoporous (a perennial wheat); 5 — triporous; 6 — poliporous (pumpkin); 7 — giant poliporous (tree-mallow — Lavatera)

Megasporogenesis or formation of female gametophyte. Ovules develop on the interior walls of ovary. Ovules are megasporangia, pented in integuments. Megasporangia are pawned in form of tubercles at placenta (place of ovule’s insertion). The main mass of ovule consists of elementary cells, developing from tubercles. These cells form nucellus (nutrient and protective tissue of embryo sac). Nucellus is covered by integuments, which leave a tight channel that is called micropyle. Part of an ovule at the opposite side to the micropyle is called chalaza.

54 The amount of ovules in the ovary is unequal (plum, cherry have 1–2; grasses have 1 ovule; poppy has many ovules), that is why the amount of seeds is unequal too. As it happens in an anther, in an ovule megasporogenesis begins from inception of archesporium (in front of micropyle). One of nucellus cells stands apart, divides by mitosis and then quaternion of megaspores is formed. Three cells of this tetrad degenerate; its inner one (the 4th cell) grows out in female gametophyte, which is called embryo sac. 4 nucleuses divide by mitosis inside the sac. As the result, eight-nucleate embryo sac is formed. Four nuclears diverge on poles; one nuclear from each pole secedes to center, forming the secondary nucleus (cytoplasm is isolated near it). At microphylous end three cells are incrusted by cytoplasm and large ovum with big nucleus and vacuole are formed (female gamete and 2 synergids). There are 3 antipode cells- on the other side. So, mature 8-nucleate sac has one ovum, 2 synergids, 3 antipodes and diploid central cell. This is the basic type of a structure natural for 82 % of plants (Angiosperms). The form of embryo sac is various: oval, straight, curved, etc. Cytoplasm of embryo sac has leucoplasts, chondriosomes, starch grains and oil drops.

Figure 45. Megasporogenesis: 1 — micropyle; 2 — integument; 3 — archegonium; 4 — primary endosperm; 5 — nucellus

Fertilization. After pollination pollen intergrowth begins. Sticky and uneven stigma’s surface helps to keep large amount of pollen grains. Besides, if partners are compatible, stigma secretes special enzyme, stimulating pollen intergrowth. Some pollen grains grow out. Success of one or another pollen depends on individual growth rate. 2 sperms and vegetative nucleus, which are located at the growing end of tube, move to it. Tube communicates with the ovule through the micropyle. Tube’s end bursts and its content erupts in embryo sac cavity. One sperm merges with haploid ovum; another one merges with diploid secondary cell. 55 Usually, only one pollen tube permeates in the embryo sac. But if there are many ovules, several of them fulfill their mission. Seed embryo develops from diploid zygote, endosperm develops from triploid one. This universal process is called double fertilization and was found by S. H. Navashin in 1898.

Figure 46. Double fertilization: 1 — anther with pollen grains; 2 — germinating pollen grain; 3 — stigma; 4 — filament; 5 — ovary; 6 — embryo sac; 7 — petal; 8 — calyx-lobe; 9 — pollen tube; 10 — vegetative nucleus; 11 — sperms; 12 — ovum; 13 — central cells; 14 — endosperm’s vestige

Distinctions of Angiosperms from Gymnosperms: 1) hybrid, triploid endosperm; 2) 2 (not 1) sperms. 3) time from pollination to fertilization is 1–2 days. Change of nuclear phases and generation alternation. Female gametophyte is an embryo sac. The male one is pollen. The ordinary type of double fertilization is called amphimixis. But sometimes an embryo develops from unfertilized elements. Apomixis is some deviations from normal course of fertilization: 1. Parthenogenesis is an embryo development from unfertilized ovum; 2. Apogamy is an embryo development from antipode or synergid; 3. Apospory is an embryo development from cell of nucellus or integument. After fertilization division of fertilized elements begins inside the embryo sac. Triploid zygote forms endosperm before embryo’s formation. An embryo develops from zygote. A perisperm can appear (from nucellus beyond the embryo sac). Integuments turn into a seed-coat. Seeds consist of an embryo, endosperm, perisperm and coat. There are 4 types of seeds: 1. The embryo occupies the whole embryo sac cavity. It displaces the whole endosperm and perisperm. Nutrients are located in seed lobes (Pod- bean family, Pumpkin family, Pink-flowered and Compound family).

56 2. The embryo occupies the minority of a seed; the larger part belongs to endosperm. There is no perisperm (Grasses, Nightshade family, Carrot family). 3. The embryo has only a perisperm. Endosperm is totally consumed by embryo while formation (Rose family, Madder family). 4. Endosperm and perisperm are preserved (pepper, lotus). While embryo formation, zygote is divided into 2 cells, forming: 1) suspensor; 2) properly embryo. Suspensor immerses embryo into endosperm and operates as a sucker. The Embryo consists of embryo rootlet, small stalk. Seed lobes are attached to the top of small stalk, forming initial plant leaves. Small stalk that is located below seed-lobes is called hypocotyl, one located above seed-lobes is called epicotyl; apex of a stem is called gemmule. Fruit is a seed with pericarp. Normal fruit is formed from an ovary: seed is formed from ovule; pericarp is from ovary’s walls. Pericarp consists of 3 layers: the outer one is called exocarp, the middle one is a mesocarp, the inner layer is called endocarp. Exocarp is a thin layer, containing 1–2 cell layer. Mesocarp is fleshy, has parenchymal cells. Endocarp is a nutshell or sap sac. Outer and inner layers can form excrescences, recesses, and hairs. They can be lignified, and its promoting spreads fruits (thistle-down, keyfruit). Pericarps are penetrated by developed system of vascular bundles. It is observed in towel-gourd fruits, in walls of poppy pod. Immature ones are rich in chloroplasts, mature pericarps are colored by anthocyanin (cherry, plum) or chromoplasts (tomato, service tree). Classification of fruits. Evolution one (according to gynoecium structure): 1. Apocarps are formed from flowers with apocarpous gynoecium and superior ovary. The most ancient apocarps are polyphyllos (crowfoot, peony), polynut, juicy apocarps (polystone-fruit). 2. Monocarps have monocarpous gynoecium, ovary is always superior. Monocarp is formed from apocarps by reduction of all fruitlets, except one. It is characteristic of Pin-flowered family, Crowfoot family: bean, monophyllos (larkspur), monostone-fruit (peach, apricot, plum); mononutlet (lady’s mantle, burnet, agrimony). 3. Coenocarps develop from flowers with coenocarpous gynoecium; their base is a compound pistil. Coenocarpous fruit can be divided by partitions, according to compound pistil partitions (bi-, tri-, multilocular fruit). Sometimes nests are totally isolated and fruit easily decays at several segments (Cabbage family). Dry coenocarps can be dehiscent, indehiscent, longitudinally separating. Juicy one usually doesn’t decay: berry, apple and others (glandular exocarp, spongy mesocarp and accrescent endocarp — citrus plants). Sometimes coenocarps devide into several parts that are called mericarps and fruit is called schizocarp. For example, Carrot family has cremocarp that devides into

57 2 mericarps during ripening. These mericarps overhang on кариофоре. Borage family and Mint family have coenobiums. These are schizocarpous fruits that consist of 4 lobes that are called eremos. 4. Pseudomonocarps outwardly look like monocaps and are formed from gynoecium, where 2 or more carpels are pawned, but only one develops. That’s why fruit is usually unilocular with 1 seedbud (nut, caryopsis, achene, sacculus). Compound fruits are usually formed from inflorescence, where fruits grow together. During the ripening they completely separate (fig, mulberry, raspberry). According to genesis fruits can be true and false. True fruits are fruits, where pericarp is formed from ovary walls. False ones are fruits, where other parts (receptacle) assist in pericarp’s formation. Conclusion. According to the State Pharmacopeia of the Republic of Belarus during fruit analysis special attention is paid to the following diagnostic features: color, smell, taste, size (length, thickness, and width), quantity of nests in fruit, character of pericarp surface, presence of conceptacle and ether-oil channels. During microscopic diagnostic attention is paid to the form and structure of epidermis cells, presence and structure features of hairs; it isimportant to notice the presence of mechanical elements in mesocarp, their shape and location, amount and distribution of ether-oil channels, vascular bundles, presence of crystal inclusions, parenchyma cell shape, etc. During analysis of seeds is considered shape, sizes, surface character, color, smell, taste; embryo shape, size and location; shape and presence of cicatricle and others. Analyzing micropreparations attention is paid to the character and structure of a seed — coat, sizes and shape of storage tissue; embryo shape and structure: seed lobes, rootlets, small stalk, gemmule. Seed coat has the most important diagnostic value. It consists of several characteristic layers: mechanical layer is formed by elongated elements like fibers of Magnolia family or Crowfoot family. The presence of slime in seed-coat epidermal cells is characteristic of several seeds. For another one is the presence of pigmented layer. Shape of endosperm cells, storage nutrients and crystal inclusion are important too.

Figure 47. Fruits: 1 — polyphylos (globe-flower); 2 — mоnophylos (brancging larkspur); 3 — polynutlet (pilewort); 4 — bean(Siberian pea tree); syncarpous: 5 — superior syncarpous pod (Saint- John’s-wort); 6 — doublewinged fruit (maple); 7 — coenobium (aromwell); 8 — inferior syncarpous pod (swordflaf); 9 — acorn (oak); 10 — nut (hazelnut); 11 — cremocarp (cow- parsnip), paracarpous; 12 — superior paracarpous pod (violet); 13 — siliele (penny-cress); 14 — inferior paracarpous pod (orchids); 15 — achene(sunflower); 16 — lysicarpous pyxidium (pimpernel)

59 LECTURE № 7 THE SYSTEM OF FLOWERING PLANTS (ANGIOSPERMS). SYSTEMATIC REVIEW OF SUBCLASSES MAGNOLIIDAE, RANUNCULIDAE AND CARYOPHYLLIDAE

The dominance of Angiosperms among the land cover determined a considerable interest of systematics to their classification. Even C. Linnaeus laid the structure of the flower down as the basis of Angiosperms’ division. The genealogical system of German botanist A. Engler on the border of XIX and XX centuries is considered to be the first one. For the first time it takes into account the possible continuity of morphological features and the evolutionary status of every genus. The largest of the immense herbaria materials in the world are located according to the system of A. Engler. Engler considered them to be unisexual, without a perianth, the flowers to be the most primitive, and taxa with such flowers — the most archaic. However, the other systematics (A. Braun, H. Gallir, Ch. Bessi, J. Hatchinson) pointed to the primitivism of large, bisexual, color polypetalous flowers and therefore considered Magnolia and Buttercup to be the most archaic. A system of Angiosperms of A. L. Takhtadzhyan that is accepted by most botanists was built according to the same system as Engler’s. In this system Magnolia are considered to be an ancient group of Angiosperm, and from their ancestors all of Angiosperms took place. The system is based on assessment of the archaic (primitive) or advanced morphological features of the plants. For this purpose, the so-called evolutionary-morphological series constructed in accordance with the criteria of evolutionary advancement is used. The initial members of each series are estimated as relatively primitive, the final ones — as evolutionary advanced. The examples of evolutionary series are: 1. Trees – shrubs – perennial grasses – annual grasses (a line usual for dicots) Herbs – secondary woody plants (for monocots). 2. Stem erect — stem creeping, curly, climbing. 3. Evergreens — deciduous plants. 4. The circular arrangement of vascular bundles — the scattered arrangement of the bundles. 5. Simple single leaves – simple dissected leaves – complex leaves – modified leaves. 6. Complex inflorescence — simple inflorescence. 7. Actinomorphic flowers — zygomorphic flowers. 8. The flowers with a large and indefinite number of parts — flowers with a small and fixed number of parts. 9. Double perianth – simple perianth – flowers without perianth. 10. Free parts of the flower — accrete parts of the flower.

60 11. Seeds of two cotyledons — seeds of one cotyledon. 12. Apocarpous fruit — syncarpous fruit. According to the Takhtadzhyan’s system, Angiosperms are divided into two classes: Dicotyledonous (Dicots) and Monocotyledonous (Monocots). Monocots took place from dicots at the early stages of Angiosperms’ evolution by accreting of two cotyledons into one. The transitional form is Nymphaeales. Dicots (Magnoliopsida, Dicotyledones) include the subclasses Magnoliids (Magnoliidae), Ranunkulids (Ranunculidae), Caryophyllids (Caryophyllidae), Gamamelidids (Hamamelididae), Dilleniids (Dilleniidae), Rosids (Rosidae), Lamiids (Lamiidae), Asterids (Asteridae). A class Monocots (Liliopsidae, Monocotyledones) includes the subclasses Alismatids (Alismatidae), Triuridids (Triurididae), Liliids (Liliidae), Aretsids (Arecidae).

CLASS DICOTS (MAGNOLIOPSIDA, DICOTYLEDONES) Subclass Magnolias — Magnoliidae — includes 5 suborders, 18 orders, 43 families, 340 genera and about 10 000 species. Modern Magnolia (remnants of a large and prosperous group of primitive flowering plants) are woody plants, rarely herbs, water plants or parasites. The conductive system is characterized by the absence of vessels, or vessels with primitive ladder perforations. There is a presence of secretory cells releasing essential oils, resins, balsams. Flowers are bisexual, spiral or spirocyclic with polymer androecium and mostly apocarpous gynoecium. Seeds contain endosperm, rarely perisperm. During our course of study, we will have three orders: Magnoliales, Laurales, Illiciales. Order Magnoliales. Includes 3 families. Representatives of the order are mostly evergreen or deciduous trees, shrubs or sometimes vines. As for anatomy they are characterized by oily or mucus cells, and the wood except tracheids has simply perforated or ladder vessels, but there are species with only tracheids. Leaves are entire or lobed, feathery. Flowers are solitary, actinomorphic, large or small in inflorescences, spiral, spirocyclic or cyclic, entomophilous. There is a single or double perianth, numerous androecium, with a primitive structure of the stamens which are often not differentiated to the staminate thread and the connective thread. Gynoecium is apocarpous, sometimes syncarpous. Seeds contain a small embryo and abundant endosperm. A characteristic feature of the flower is a proliferating receptacle, especially growing in fruit. The centerpiece of the order takes a family Magnolia. Family Magnolia — Magnoliaceae. Includes 12 genera and about 240 species, mostly spread among the mountain forests of south-East Asia and the Southwest of North America. One of the representatives of the family — Magnolia obovate (Magnolia obovata L.) is found in natural form on Kunashir island (Kuril Islands), and the Magnolia grandiflora (Magnolia grandiflora L.) is

61 grown in the gardens and parks of the Black Sea coast. There also grows a tulip tree (Liriodendron tulipifera L.). Magnolia are deciduous or evergreen shrubs or woody plants. Leaves are simple, alternate, with large caducous stipules. The leaves contain oil glands. Perianth is 3–6–9-membered, single or double, its painted segments are arranged cyclically in the 1–2 range. The stamens have a petaloid form and are numerous. Gynoecium is apocarpous, often closed carpels are arranged on a conical receptacle spirally. *Са3Со3+3А∞G∞ Magnolia flowers are adapted to pollination by beetles, that penetrate the unclosed buds. Fruits — spiral leaflet, polynutlet. The seeds are surrounded by lush brightly colored cover and suspended on threadlike funicles, spread by birds. Winged nuts of tulip tree are spread by wind. Representatives — Magnolia grandiflora L., Liriodendron tulipifera L. are cultivated as ornamental plants. Magnolia grandiflora L. — a tall tree with showy large evergreen leaves with stipules accrete, which cover the gemmas. In the leaf’s mesophyll there are large glands with ethereal oil. Flowers are large, 35 cm in diameter, tepals are white with a slightly yellowish tinge, and fragrant. Their number is uncertain, but one can see the upper 3 tepals forming a circle, for there is a tendency to form a triple perianth. Androecium is numerous, ribbon-like stamens are not differentiated on the thread, and staminate connective, anthers are arranged in long strips along the edges of the stamens and above the anthers acts on well- marked connection. Pollen falls on the spoon tepals and is eaten by beetles — pollinators of magnolias (kantarofiliya). Gynoecium is represented by numerous free carpels, arranged in a spiral, with sessile stigma. Preparations from the leaves of Magnolia grandiflora are used in the treatment of hypertension. Order Laurales. Contains 11 families. The deciduous and evergreen trees and shrubs belong to this order, rarely shrubs, and even rarer herbs with alternate or opposite leaves. Flowers are cyclical, mostly bisexual, sometimes unisexual, small, gathered in clusters. A characteristic feature of the flower is an extended and a concave receptacle and floral tube (accrete leaves at the base and stamens adnate to it). Gynoecium is apocarpous, often monomer (of one carpel), sometimes syncarpous. The most famous families are the Monomiaceae and the Lauraceae. Laurel family — Lauraceae. It includes up to 45 genera and 2500–3000 species, distributed mainly in tropical and subtropical areas. Almost all members of the family are trees and bushes. The leaves are alternate, simple, whole and leathery. Wood, bark and leaves contain essential oil containers, so some of them are used as spice plants, but there are poisonous ones among them, giving the poison curare. Flowers are actinomorphic, bisexual or dioecious, small, inconspicuous, collected in (paniculate or racemose) inflorescences. Perianth is

62 simple, 4–6-membered, in poorly differentiated circles. The stamens are arranged in four circles (anthers have 2–4 valves, depending on the number of slots). The stamens located in the inner circle turned into nectaries and staminodes. Gynoecium is a monomer, seeded bacciform juicy fruit is formed of it, surrounded at the base with the peduncle grounds (cupula). Ovary is superior, rarely lower. * P3+3А3+3+3+3G(3)– Representative of Lauraceae is laurus noble (Laurus nobilis L.), cultivated on the Black Sea coast of the Caucasus. Dry leaves of this plant, or laureate leaves, are used as spices. Cinnamomum Ceylon (Cinnamomum zeylanicum Blume) — the bark is used as a spice, Ceylon cinnamon. Cinnamomum camphor (C. camphora (L.) J. Presl) is a source of natural camphor. Avocado (Persea americana) is a fruit tree. The fruits contain up to 32 % digestible oils and are a valuable dietary product. Order Star Anises — Illiciales. Wood trees, shrubs and vines belong to this order along with evergreen or deciduous. Leaves are without stipules. The wood has primitive vessels with ladder perforations. Flowers are bisexual or unisexual, spiral or spirocyclic. The order consists of two families with a small amount of genera. Family Star Anises — Illiciaceae. The monotypic family containing one genus Illicium — star anise, which includes about 40 species. These are evergreen low trees or shrubs with simple alternate leaves and single flowers. The most famous is star anise (I. verum Hook. F.), which is used as a spice. Fruits contain anethole and therefore smell like anise from family Umbelliferae. Another representative of this family is a sacred star anise (I. anisatum Hook. F.), which has poisonous fruits. It is bred as an ornamental plant at Buddhist temples, bark and seeds are used as incense in the East. Family Lemongrass — Schisandraceae. It contains two kinds: Lemongrass — Schisandra sp., kadsura — Kadsura sp. This is a deciduous or evergreen vine with simple, alternate leaves. The wood has vessels with primitive segments that have a ladder perforation. The flowers are not large, but rather axillary with strongly elongate (like lemongrass) receptacle; the fruits are juicy indehiscent polyleaflet. Lemongrass chinease is widely spread (Schisandra chinensis (Turcz.) Baill.) — a major climber far 4–15 m in length. Leaves are simple, small, discharged for the winter. The flowers are white or pink, fragrant, waxy, monoecious, monosexual, with long reddish stalks. Perianth consists of an indefinite number of petals, stamens in the male flowers are of 1–3, carpels in female flowers are of 30–40. Sometimes there are bisexual flowers. Typically, lemongrass is bioecious, but sometimes monoecious. The fruit is a juicy leaflet. The skin and flesh of lemongrass are sweet-sour with a lemon smell and the whole fruit with seeds is salty. Lemongrass is widely used as a medicinal,

63 ornamental and food plant. The fruits and seeds contain citric acid (10–11 %), malic acid (7–8 %) and other acids, and essential fatty oils, tannins, monocarbohydrates, proteins. In addition to essential oil they contain lignans, determining the tonic effect of fruit, from which the drug is widely used in traditional medicine. When rubbing the stems, leaves and roots it smells like lemon. Subclass Buttercups — Ranunculidae. Subclass combines 4 orders and 13 families, 200 genera and 4000 species. Representatives of the subclass on a number of grounds are close to the magnolia subclass, but are more highly developed. Grass plants with a developed dominate conduction system and secretory cells are rare. Flowers are bisexual, spiral or spirocyclic, with numerous androecium, the gynoecium is apocarpous in many cases, but in many evolutionary lines of the subclass a syncarpous gynoecium is observed. Seeds contain endosperm and a small embryo. The most common in the temperate zone are representatives of the orders Buttercups, Poppy flowers. Order Buttercups — Ranunculales (Ranales). Includes 8 families. Grass plants are dominant, shrubs and vines are also sometimes found. While widely distributed in the temperate zone, they are aslo found in tropical and subtropical zones. Segments of vessels have simple perforations. Secretory cells accumulate alkaloids. Leaves are simple or compound, without stipules. Flowers are bisexual or unisexual, actinomorphic (with an indefinite number of petals arranged in a spiral) or zygomorphic. Perianth is simple or double, or non- existent. Numerous androecium, gynoecium is apocarpous, seeds contain copious endosperm. Buttercup family — Ranunculaceae. It has 60 genera and over 2000 species, distributed mostly in the temperate and cold regions of the globe. The vast majority of members of the family are grass perennial (rarely annual) plants, often living in damp places, there are shrubs and vines. Leaves are simple, dissected, alternate. The structure of the flowers has sufficient diversity: the flowers have often a simple uncertain or 3-membered perianth, but some genera have double perianth, either indefinite or of a five-membered structural plan. The flowers are nearly always bisexual, actinomorphic, excluding larkspur and a fighter which are zygomorphic. Calyx is of 2–6, usually 5 colored sepals, their number is not constant. The petals may be reduced or partially converted into staminodes. Androecium is numerous, stamens arranged spirally. Gynoecium is apocarpous, occasionally monokarpous (black cohosh). *Са5Со5А∞G∞ The flowers are pollinated by insects. Flowers are characterized by the presence of nectaries of various types: a pit at the base of the leaves, or modified stamens or petals. Fruits are polyleaflets, polynutlets. Seeds pertains in an endozoohoric way. Vegetative organs contain alkaloids and are often toxic.

64 Among the representatives are widely spread plants, used in traditional medicine, fancy ones. Typical representatives of the family are: Adonis vernal (Adonis vernalis L.), marsh marigold (Caltha palustris L.), Anemone nemorosa (Anemone nemorosa L.), European globe (Trollius europaeus L.), meadow buttercup (Ranunculus acris L.), high larkspur (Delphinium elatum L.), monk blue (Aconitum napellus L.), Aquilegia vulgar (Aquilegia vulgaris L.). The marsh marigold, anemones nemorosa and European globe are characterized by a simple perianth, all the other representatives have a double perianth. Buttercups are rich in substances of secondary metabolism. They are found of glycosides, saponins (Adonis varnal), alkaloids (monk blue, larkspur high). Most members of the family are poisonous. Family Barberries — Berberidaceae. Includes 14 genera and 650 species that are distributed in temperate and subtropical latitudes. This family is represented by shrubs and perennial grasses. Leaves are simple or complex, some modified into spines. The flowers are small, actinomorphic, bisexual, two- , rarely threemerous. Numerous stamens are arranged in two circles, gynoecium composed of one carpel with one or many ovules. *P3x4 (6 internal leaves turned into nectaries) A3+3G1. Its fruit is often bacciform, but sometimes a capsule. Barberries contain alkaloids. A typical representative of this family is the common barberry (Berberis vulgaris L.). It is a richly ornamental flowering shrub that has thorny shoots. Leaves are simple or complex, some of them modified into spines. The flowers are small actinomorphic, collected in inflorescence of brush. Petals have nectaries at the base. Fruitage is represented by berries. The representatives of this family contain alkaloids. Alkaloids, primarily berberine, which is used in obstetric practice. The family of Peony — Paeoniaceae. This family includes a single genus of peony (Paeonia) with 35–40 species. There are both grass and shrub, the so- called ―tree‖ peonies. Flowers are terminal, large, single, bisexual, actinomorphic. Perianth consists of 5 hard sepals and 5 (10–12) of large brightly colored petals. Stamens are numerous, gynoecium is apocarpous of 2–5 carpels. *Ca5Co5A∞G1-8. Fruit is polyleaflet. Representatives: Marin roots (peony) (Paeonia anomala L.), fine-leaved peony (P. tenuifolia), white-flowered peony (P. lactiflora Pall.). Peonies are valuable medical plants, roots of Marina root are used as a sedative tincture. On the basis of wild varieties there is created a large number of ornamental ones. Poppy family — Papaveraceae combines 24 kinds and 250 species distributed in subtropical and temperate latitude. These are perennial or annual grass plants, sometimes half-shrubs or shrubs with alternate, simple, without stipules leaves. They are characterized by presence of lactifers with yellow, orange or colorless juice, which contains alkaloids. The flowers are bright, large,

65 actinomorphic, solitary or collected in inflorescence. The calyx contains two sepals that fall down by the time the flower is uncovered. Sometimes, there are more than two sepals (three or four). There are four petals, arranged in two circles. Androecium is numerous, rarely four, which are arranged in a spiral. Gynoecium is parakarpous of many or 2 carpels with the upper, sometimes semi-inferior ovary.*Ca2Co4A∞G(2). Fruitage is represented by capsules. Seeds contain an abundant endosperm. Poppy hypnotic, opium (Papaver somniferum L.). It is an annual plant with simple coarsely jagged or incised-lobed leaves. The lower leaves are on short petioles, the upper — surround the steam. On the underside of the leaf blade there are sparse hairs. Flowers are regular, bisexual. Perianth is doubled, 2 sepals (caducous), 4 petals (arranged in two circles). Androecium is numerous (8–20), the stamens thicken up. The gynoecium is formed with a spherical pestle with stellate hairy stigma. Fruitage is represented by capsules. Seeds are small, contain alkaloids. They extract lactic juice — opium from green bolls. Like a weed there exists cornrose (P. rhoeas L.). Celandine, (Chelidonium majus L.) — a perennial plant with simple pinnatisect, with ear-shaped slices clauses at the base leaves. The upper side of the leaves is green, the bottom is glaucous. There are lactifers. Flowers are regular, bisexual. Perianth is double, of 2 sepals and 4 petals (arranged in two circles). Androecium is numerous. The gynoecium consists of two carpels. Fruitage is represented by a pod or capsule. The seeds contain a large white aril. The plant is poisonous, all organs contain alkaloids. Orange juice of celandine is used for skin diseases. As medical plants in the SF RB are leaves of maclay are listed as an antimicrobial remedy (Macleaya cordata). Family Fumitorious — Fumariaceae is a small family. The grasses with a bad cut, alternate leaves, without latex, but with secretory cells. The flowers are small, zygomorphic, containing spur with a nectar, there are 2 sepals (usually falling); and four petals. Androecium consists of hardly constructed 2 stamens. Each stamen has 3 anthers, 2 of which are bilocular and the middle one is tetraocular, filaments are accreted at the base. The gynoecium contains 2 carpels. Fruit is represented by silicular capsule or nutlet. Fumaria medical (Fumaria officinalis L.) is an annual plant with leaves repeatedly dissected into numerous small lobes. The flowers are small, collected in an inflorescence brush. Fruit is represented by an indehiscent nut. The herb is used as a diuretic and lactiferous tool. Corydalis coriaceous (Corydalis solida) — a characteristic feature is the presence of color spur formed by petals. Subclass Cloves — Caryophyllidae. Subclass combines 3 orders, 19 families, 650 genera and 11500 species. They are dominated by grass plants, shrubs, bushes, occasionally there are medium-sized tree forms. There are vessels with simple perforations. Leaves are simple, entire. Flowers are bisexual,

66 rarely unisexual, actinomorphic, occasionally without petals. Androecium is numerous. Gynoecium is syncarpous, occasionally apocarpous. Seeds contain perisperm and bend embryo. Order Cloves — Caryophyllales comprises 17 families, the most famous are cloves and goosefoot, cactus, amaranth. Grass plants, rarely shrubs. Leaves are entire, opposite. Flowers are bisexual or unisexual, actinomorphic, 5-membered with a double perianth. Androecium is numerous or well-defined, some stamens are arranged in two circles. Gynoecium is lysicarpous. Fruit is represented by a capsule. Family Cloves — Caryophyllaceae has 80 genera and 2000 species. Most are annual and perennial grass plants, shrubs are also found, ubiquitously. Leaves are opposite, rarely alternate, but with membranous stipules. Flowers are solitary or collected in dichasial inflorescences. Actinomorphic and is 5-membered with a double perianth. Sepals are free or accreted into a tube, petals are free or accreted. Some species have a narrow base of petals, or claw, that has a wide return flange (repeatedly dissected into lobes) the tip, or limb. At the point of transition into the nail petaloid or membranous outgrowths — appendages — are visible, there exists a paracorolla. Stamens are from 4–5 to 10, 2–5 carpel gynoecium is syncarpous turning into lysicarpous, ovary is superior. *Са(5) or 5Со5А5+5G(2-5) Fruit is represented by a capsule, nutletkind or less berries. Seeds have fleshy appendages and are carried by ants. The representatives of this family are: – Wood stitchwort (Stellaria nemorum L.) — a perennial herb with a recumbent, rising stem with adventitious roots. The leaves are heart-shaped, softly hairy, petals are deeply incised into 2 parts; – London pride (Saponaria officinalis L) — a perennial herb with erect stem and broadly, sessile leaves. Petals have paracorolla, dissected along the edge into shares. The roots contain saponins and are used as expectorants, a diaphoretic and antirheumatic remedy in metabolic disorders; – Meadow pink (Dianthus deltoides L.) — a perennial herb with erect stem and narrow-lanceolate, sessile, rough-hairy leaves. Calyx has bracts membranous scales; – Cockweed (Agrostemma githago L.) — perennial herb. Stem is branching. Leaves are linear-lanceolate with the hairs. Flowers are solitary. All these representatives have the fruitage of capsule. Many ornamental plants are among the representatives of this family. Family Goosefoot — Chenopodiaceae has about 105 genera and 1600 herbaceous and shrub species, common everywhere, but especially spread on saline soils (Haloxylon, Salicornia, Solenokolosniki, Soups). The flowers are small, actinomorphic, bisexual or unisexual, without petals and are usually collected in inflorescence brush or whisk. Calyx has five nondescript greenish

67 leaves and is surrounded by large bracts. Perianth can be completely reduced. Stamens are 5, rarely 1–4. Syncarpous gynoecium consists of 2 (3–4) carpels forming a top or semi-inferior ovary with free or almost accreted columns. *P(5)А5G(2-5) Fruit is one-seeded, indehiscent, nut-kind, falling along with the cup, the lobes of which grow in the wing or hooked growths. Sometimes a cup becomes fleshy and fruit looks like juicy berries (Chenopodium foliosum L.). In the formation of the fruits (in some cases) bracts take the engagement (Atriplex sp.). When accretion of sepals and bracts, such as beets and spinach, multiple fruit is formed. Representatives of weeds are characterized by heteromorphism of flowers, fruits and seeds within even a single inflorescence. The flowers are wind- pollinated, the spread of seeds is based on the principle of a rolling stone — field. The representatives of this family have nutritional value. Spinach garden (Spinacia oleraceae), beets (Beta vulgaris L.) contain large amounts of vitamins, proteins and mineral salts. As a substitute for spinach you can use the young shoots of wild saltbush (Atriplex sp.) or goosefoot (Chenopodium sp.). Cactus family — Cactaceae includes 105 genera and 2200 species distributed in extremely arid areas. Most of the species lose its leaves, their cylindrical or spherical fleshy stalk (performing water-saving and photosynthetic function) that carries bundles of thorns, covered with hairs, which are modified kidney scales. The representatives of this family are used for decorative purposes (Zigocactus, Epiphillum, Echinopsis, etc.). Amaranth family — Amaranthaceae has 65 genera and 850 species distributed mainly in tropical areas. Amaranth is an annual or perennial grass with entire alternate leaves. The flowers are small, actinomorphic, bisexual, collected in inflorescence brush or broom. Corolla is calyx-kind (no petals), the calyx consists of 5 filmy scales and surrounded by bracts that may remain after flowering, so it seems like an unfading flower. 5 stamens, gynoecium consists of 2–3 syncarpous carpels forming the top unilocular ovary. Fruit is represented by a nut, berry or rare pyxis. erwa wooly or half-floor are used in medicine (Aerva lanata (L.) Juss.) to treat urolithiasis. Most of the representatives have ornamental value: amaranth-bleeding (Amaranthus caudatus L.), celosia (Celosia argentea var. Cristata). Leaves and shoots of some Amaranths are used for food (A. caudatus L., A. Cruentus L.). Order Buckwheat — Polygonales includes only one family of buckwheat (Polygonaceae) witgh 30–35 genera and over 1000 species. Representatives of the family are widespread, are mainly found in temperate regions. Buckwheat of temperate regions is an annual or perennial grass. Its characteristic feature is the presence of the socket (accrete in a tube stipules), covering the stem. The flowers are small, actinomorphic, bisexual (rarely unisexual), collected in terminal racemose inflorescence. A simple perianth which consists of three,

68 rarely two-five greenish or colored sepals. Stamens are of 3–9, nectaries are located between their bases. Lysicarpous gynoecium consists of 3, rarely 2–4 carpels, forming a unilocular ovary, having one ovule and free or accreted bars. *P3-6A5-9G(3) or (2-4). Fruit is represented by a nutlet with a number of faces which equal the number of carpels. Buckwheat is a valuable food, industrial and medicinal plant. The leaves of sorrel (Rumex confertus L.), are stalks of rhubarb (Rheum rhaponticum L.). They contain organic acids and vitamins that are used for food. It is widely used as a medicinal plant different kinds of highlanders: Highlander pepper (Polygonum hydropiper L.), highlander maculosa (Polygonum persicaria L.) used as a hemostatic agent in uterine and hemorraidal bleedings. Bird highlander (Polygonum aviculare L.) is used as an anti-inflammatory agent, as well as means of facilitating discharge of stones in the kidneys and bladder. The rhizomes of snake highlander (Polygonum bistorta L.) are used as an astringent in inflammatory diseases of the mucous membranes in the mouth. Buckwheat (Fagopyrum esculentum Moench) can be used for industrial production of routine.

69 LECTURE № 8 SYSTEMATIC REVIEW OF SUBCLASSES HAMAMELIDIDAE, DILLENIIDАE

Subclass Gamamelidids — Hamamelididae. Includes 16 orders, 22 families, 71 genera, 1500 species. It is an ancient group of flowering plants that evolved from Magnoliids. Avascular forms are reserved. All the members of this division are woody plants, In their evolution a simplification of the flower (the perianth is reduced) was observed. The flowers are unisexual. There is a transition to anemophily. Carpels in some cases remain free. Order Beeches — Fagales. This order includes one of the Beeches’ family, so characteristics of the order coincides with the characteristics of the family. Family Beeches — Fagaceae. Includes 8 genera and more than 900 species distributed throughout, except for tropical and polar regions. Beeches are large deciduous or evergreen trees, rarely shrubs or bushes. Leaves are simple, alternate, leathery, with caducous stipules. The flowers are unisexual, in ament or glomus tirses where the partial inflorescence represent a dichasia, sometimes reduced to a single flower. Some female dichasia are surrounded by scaly, lumpy, or a prickly cupped wrapper or cupule, which is formed by the sterile branches of florets. Perianth is simple, it consists of 4–7 lobes, is plain, accreted at the base of the male flowers, and adherent to the ovary in females. Stamens are free, Its number is twice as high as the number of sepals. Gynoecium is syncarpous of 3 carpels forming a 3-alopecia ovary. Each dissepiment has 2 ovules from which Only one develops. The ovary is inferior. Fruit is represented by a seeded nut, completely or partially surrounded by woody cupule called acorn. Representatives: Castanea sativa Mill. — real chestnut, fruits are used in food, Fagus orientalis Lipsky — Eastern beech, fruits contain edible oil. Quercus robur L. — old or pedunculate oak, bark and leaves contain tannins. ♂*P(5-9)A(6-9)G0, ♀*P3+3A0G(3) Quercus infectoria Oliv. — oak dyeing, galls formed on the leaves are a source of tannin. Quercus suber L. — cork oak, a source of stoppers. The order Birches — Betulales. One family — Betulaceae relates to order, and includes 6 genera and 150 species. The family is represented by deciduous trees and shrubs, forming small-leaved forests. Leaves are simple, entire, alternate, with deciduous stipules, usually pubescent. The plants are dioecious. The flowers are small, inconspicuous, unisexual and collect into an inflorescence. Plants are monoecious. Male flowers are in drooping ament inflorescences, female — in sticking pineal, which are based on dichasium. Bracts of female flowers often

70 grow together, forming a coating of dichasia scales. In other cases, for example in hazel, leathery scales form cupule, covering the fruit. Perianth is reduced. Male flowers have from 2–4 to 12 and many stamens. ♂*P2A2G0. Female have pseudomonocarpous gynoecium consisting of two carpels forming a unilocular ovary with long stigmas. ♀*P0A0G(2). The flowers are pollinated by the wind. Fruit is represented by a nut or nutlet, with membranous wing-like outgrowths on the sides. Representatives: Alnus glutinisa (L.) Gaerth. — sticky alder. ♂*P4A4G0, ♀*P0A0G(2) A. incana (L.) Moench — grey alder. Betula pendula Roth — white birch. ♂*P2A2G0, ♀*P0A0G(2) B. pubescens Ehrh. — velvety birch. B. nana L. — dwarf birch. Corylus avellana L. — vulgar hazel. The buds and leaves of birch trees are used in medicine as a bactericidal and cholagogue. Order Walnuts — Juglandales. This order involves a single family walnut — Juglandaceae, which includes 7 genera and 60 species. Walnut is a deciduous tree with large pinnate leaves with aromatic glands. Plants are monoecious. The flowers are unisexual, small inconspicuous, clustered in unisexual inflorescences — male in ament tirses ♂*P2,4A8-10G0, while female are in the same tirses, only in groups or single. The basis of the inflorescences contains dichasia, reduced to a single flower. ♀*P4A0G(2). A perianth is absent or greatly reduced. 2–100 stamens, gynoecium of two carpels, pseudomonocarpous. Fruit is represented by a drupe with leathery exocarp. A representative of this family is the greek walnut (Juglans regia L.), the fruits of which are used in. It has valuable timber. Subclass Dilleniidae. Phylogenetically Dilleniidae represent one of the main groups of flowering plants. This class includes 31 orders, 97 families, around 1910 genera and about 36 000 species. In the most primitive Dilleniidae more commonalities with the magnolia subclass were preserved. This subclass belongs to the temperate zone plants, as well as representatives of the tropical and subtropical flora. Life forms are represented by trees, shrubs and grasses. The conduction system is characterized by the presence of vessels with a ladder and a simple perforation. Flowers are bisexual, or unisexual, with a double apetalous perianth or for more primitive flowers with a spirocyclic perianth. The members of this family have a numerous androecium. The gynoecium for primitive flowers is apocarpous, more advanced flowers have a syncarpous one; seeds contain endosperm. In this course we will get acquainted with orders Theales, Violales, Malvales, Ericales, Primulales, Cucurbitales, Urticales. Order Theales. Includes 12 families and is represented by woody plants, shrubs, vines, and — very rarely — herbs. Leaves are simple, alternate. Flowers

71 are actinomorphic, rarely zygomorphic and bisexual. Androecium contains accreted stamen filaments. Gynoecium is syncarpous or secondary apocarpous, ovary is superior, endosperm is often with seeds. The fruit is represented by a capsule, drupe, berry. Most plants of this order are spread in subtropical and tropical zones. In the temperate zone this order is represented by families of tea and John’s-worts. Tea family (Theaceae) includes 24 genera and about 560 species. All representatives are evergreen trees or shrubs. Leaves are simple, alternate, leathery. Flowers are single, often large, actinomorphic, bisexual. Calyx has 5–7 sepals, corolla is usually 5 — membered, sometimes with 4–9 petals. Androecium is numerous of growing together or free stamens, occasionally 5–10. The gynoecium is of 2–3–5 or more carpels, syncarpous. The ovary is superior, fruit is represented by a capsule, a dry drupe, or berry. Representatives tea bush (Thea sinensis L.) — a shrub with simple thick, leathery, alternate leaves, Ca5-6Co5-9А∞G(3-5), a fruit is represented by a capsule. It contains vitamins, tannins, caffeine, alkaloids. Ornamental plants are represented by the Japanese camellia (Camellia japonica L.), sasanqua (C. sasanqua L.) having the largest terry white, pink and red flowers. Family John’s-worts (Hypericaceae) contains 9 genera and about 400 species. Along with wood and bush forms, grasses are also dominant. Leaves are simple, opposite with point translucent veins. Flowers are solitary (like tea flowers) or collected in cympid inflorescences. Stamens have long filaments, accreted in beams. Gynoecium is syncarpous, ovary is superior with free columns Ca(5)Co5А∞G(3), a fruit is represented by a capsule. Representatives: St. John’s-wort (Hypericum perforatum L.), St. John’- wort speckled (H. maculatum L.) — meadow grass used in folk and scientific medicine as anti-inflammatory, antidepressant and astringen remedyt. Order Violales unites 15 families and is represented by trees, shrubs and grasses. Leaves are alternate or opposite with stipules. Flowers are actinomorphic, zygomorphic, bisexual. Androecium is numerous or of 5 stamens with free or accreted stamen filaments or anthers. Gynoecium is paracarpous, ovary is superior, semi-inferior or lower. The fruits are capsules, berries or nuts. The plants of this order are widely abound. The representatives of the family Violaceae are of great importance. Violet family (Violaceae) is represented by 29 genera and 900 species. A lot of grasses, shrubs, vines, large trees belong to this family. Leaves are alternate or opposite with stipules. Flowers have a double perianth, actino- or zygomorphic, 5-membered. Zygomorphic flowers have one petal with a larger bag-shaped spur at the base. 5 stamens may coalesce into a tube surrounding the ovary. Syncarpous gynoecium consists of 3 carpels, forming the upper ovary, ↑ Ca5Co5А5G(3). Fruit is the box (3-leaf), rarely a berry or a nut.

72 Representatives: Violet three-colored (Viola tricolor L.), field violet (V. arvensis L.), the grass is part of the pectoral and diuretic fees. Violet amazing (V. mirabilis L.) and others. Order Heathers — Ericales. This order is closely associated with the order Theales. It is dominated by shrubs and trees, only occasionally is marked by saprophytic perennial grasses and plants. Members of this order are common in different areas and different environmental conditions. Leaves are alternate, rarely opposite or whorled. The flowers are mostly bisexual, actinomorphic, gamopetalous, sometimes free-petalous, stamens and petals are attached to the side of the hypogynous nectar disc, and the stamens are opened up with apical pores. Parakarpous gynoecium is of three or more carpels, the ovary is superior or inferior, fruits are represented by berries, capsules and drupes. Heathers family (Ericaceae) — the largest in the order Ericales, is very typical for the tropical and subtropical areas, but some of its representatives are characterized by plants of temperate and arctic zones. More often they are shrubs or woody plants, sometimes shrubs or vines, evergreen or deciduous, with alternate leaves. Flowers are 4–5-membered, often collected in racemose, umbellate, corymbose inflorescences, bisexual, actinomorphic, often with accreted cup and accreted whisk. Usually there are 4–5 sepals, sometimes 8, the same number of petals, often sympetalous, there are twice as many stamens as petals, they are arranged in two circles or less, they are as much as the petals, sometimes up to 20. Gynoecium is of 4–5, sometimes 10 carpels, the ovary is superior, Ca(4-5)Co(4-5) or 4-5А8,10G(4-5). Fruit is represented by a box or a berry. Representatives: wild rosemary (Ledum palustre L., shoots contain essential oils and are used for diseases of the respiratory tract; ling (Calluna vulgaris (L.) Hill), flowers and sprouts are used in traditional medicine, common bearberry (Arctostaphyllos uva-ursi (L.) Spreng.), leaves and shoots are used for diseases of the urinary tract. The family of cranberries (Vacciniaceae) combines bushes, shrubs with evergreen or deciduous leaves. Cranberries differ from heathers only by the presence of an inferior ovary. Flowers are 4–5-membered, usually sympetalous, androecium of stamens, the number of which is twice higher than the number of shares of perianth, gynoecium has 4–5 carpels, fruitage is represented by a berry. Representatives: cow berry (Vaccinium vitis-idaea L.), leaves and shoots are used as a disinfectant and mild diuretic; blueberry common (Vaccinium myrtillus L.), the fruits are used in homeopathy; the fruits of swamp blueberry are used (Vaccinium uliginosum L.); fenberry (Oxycoccus palustris Adans.), the fruits of which are used as an antipyretic and there are anti-inflammatory agents.

73 Order Primulas — Primulales includes 4 families represented by trees, shrubs and grasses with alternate or opposite, simple (sometimes whorled) leaves without stipules. Flowers are bisexual or unisexual, actinomorphic, usually 5-membered. Sepals fare free or accreted, corolla is sympetalous, stamens are arranged in one or two rounds, adnate to corolla, gynoecium is lizycarpous, ovary is superior or inferior, sometimes semi-inferior, Ca(5)Co(5)А5G(5)–. The fruit is represented by berries, drupes and capsules. Primulas family (Primulaceae) consists of 30 genera and 1000 species. This is the largest family in the order Primulales, very widespread but dominant in the temperate and cold regions of the northern hemisphere. Many grasses belong to this family: water grasses, shrubs. Leaves typically form a rosette, are alternate, opposite and whorled, simple or lobed, sometimes strongly dissected. Flowers in a variety of inflorescences are sometimes solitary, sometimes 5-, sometimes 4–6–7–9-membered. Calyx is always accreted, corolla is also gamopetalous (petals can coalesce only at the bottom), androecium has the same number of stamens as petals, gynoecium has 5 carpels, ovary is superior or semi-inferior. Flowers are characterized by heterostylism. Representatives are: cowslip primrose (Primula veris L.), leaves and roots are used in homeopathy; decorative plants — bear’s-ear (P. farinose L.), European starflower (Trientalis europea L.). Order Mallows — Malvales includes 11 families. This order includes trees, shrubs and grasses, with alternate simple or complex leaves with stipules. It is characterized by slime cells or channels, as well as abundant pubescence, especially on fruits and seeds. The flowers are usually bisexual, actinomorphic, usually 5-membered; stamens are arranged in two circles, the outer sometimes turns into staminodes and the inner split into a large number of stamens accreted into beams or columns with their yarns. Gynoecium is often parakarpous, but usually secondary syncarpous, the ovary is superior. The fruits are dry, drop, such as capsules, or fractional. The family of Teils (Tiliaceae) includes 46 genera, 450 species. It consists mainly of tropical and subtropical plants: deciduous trees, shrubs, sometimes semi-shrubs and grasses. However, some members are in temperate climates. The leaves are alternate, simple, with deciduous stipules. Flowers are bisexual, actinomorphic, 5-membered, rarely 3–4-membered, sepals and petals are of the same number, androecium is numerous, with stamen filaments, accreted into bundles, or only at the base, or free. Gynoecium is of two or more carpels, paracarpous, but as a result of fusion of placentas in the center of the ovary, became a multi-cavity. Ca5Co5А5 or ∞G(5). Stamens are accreted into five beams. The flowers are collected in inflorescence. The fruits are dry and juicy. A typical representative of this family is the heart-shaped teil (Tilia cordata L.), inflorescences (teil’s blossom) is used as a diaphoretic.

74 Family Mallows (Malvaceae) includes 85 genera, 1000 species. Members of this family are widely distributed around the globe and dominated in the tropics, but a number of its representatives are and in the temperate zone. This family includes grasses, shrubs and trees with simple, alternate leaves with stipules. Flowers are actinomorphic, bisexual, 5-membered. Calyx is of 5 sepals, often with epicalyx, petals come in five, free or accreted; androecium is numerous in 2 large circles accreted into a tube with the stamen filaments, gynoecium is of 2–5-many carpels. Ca3 or (3)Co5А(∞)G(∞). The flowers are usually collected in inflorescence. Fruit is a capsule, or decays into mericarps (fractional), sometimes berry or key. Characterized by hairs on the seeds or in the fruits, mucose channels, and outline of the phloem of the stem in cross- section resembles that of the family Tiliaceae — it has a trapezoidal shape. Representative of this family is the medicinal marsh-mallow (Althaea officinalis L.), the roots of which contain up to 35 % of mucus and starch. Along with this kind of marshmallow armenian marsh-mallow is used (A. armeniaca Ten.). Order Capers — Capparales. Woody plants, shrubs, and numerous grasses with simple, alternate leaves which are widely distributed around the globe belong to this order. Flowers are actinomorphic, bisexual, with a double perianth, 4–5-membered. Numerous androecium of 6 stamens, gynoecium is parakarpous of 2–4-many carpels, ovary is superior. Fruit is represented by a capsule, silicular boxes, pods, berries. Family Cruciferous (Brassicaceae or Cruciferae) — the largest family among Capparales, which are spread mainly in the northern hemisphere. Most cruciferous are grasses, rarely half-shrubs, even more rarely shrubs, with alternate leaves without stipules, often forming a rosette. Flowers are actinomorphic with a clear form of the structure: four sepals, four petals with a visible marigold, androecium consists of six stamens in two rounds, 4 longer and 2 shorter, sometimes 4, 2 or 16, gynoecium is of two carpels, ovary is superior, Ca4Co4А4+2G(2). The flowers are small, strongly lengthening racemes, while the blossom and fruit ripe. Fruit is represented by a pod, silicle or nutlet. Breeds of gillyflower (Matthiola annua sp.), having a bright, often double flowers with a strong aroma are commonly known among the most popular ornamental plants. Large flowers of gillyflowers show cruciferous flower shape. Cole seed (Brasssica campestris L.), is an annual plant with a rosette of lyrate bluish leaves, yellow flowers and a large, typical for cruciferous, long pods. The genus Brassica includes many crop plants: different breeds of garden cole (B. oleracea L.), swede (B. napus L.), turnip (B. rapa L.) and others. Medical plants include a shepherd’s purse (Capsella bursa pastoris (L.) Medik.); the herb is used to treat liver diseases, and uterine bleeding. Order Osiers — Salicales. Osier family (Salicaceae). This family consists of trees, shrubs and small shrubs. It also contains 3 genera: Salix — osier, 75 Populus — poplar and Chosenia — chosenia and 400 species distributed in temperate, arctic areas, and in the areas of high mountains. All osiers have simple alternate leaves with stipules and small unisexual flowers, gathered in ament clusters. The flowers are deprived of the perianth. Androecium can be numerous or in 2–3–5 stamens, gynoecium is parakarpous, fruit is represented by a capsule, reversed with 2 doors. The genus Populus — poplar, wind-pollinated plant, dioecious. Poplar is blooming in the spring, before leafing. At this time on male examplars appear inflorescence aments, drooping, with a flexible axis, bearing brownish and tight- jagged on the edge bracts; male flowers develop in the axils having numerous stamens with purple anthers. They have no perianth. Female examplers develop similar inflorescences, their axises also hold bracts. Gynoecia of polars are located in in their axle on the wineglass disks, which contain a massive stigma. Ripe fruits are medium-sized capsules, dropped-down with two wings, producing numerous small seeds, wrapped in fine hairs. At the time of mass ripening Poplar abundantly ―powders‖, bringing inconvenience to citizens. That is why only male specimen of poplars should be propagated. Representatives of this genus include: poplar fragrant, (Populus suaveolens L.), a plant with large broadly-egg leaves, a strong flavor, thanks to the resins covering the surface of the bud scales; black poplar (P. nigra L.), whose buds are used as an antirheumatic agent. Poplars are highly phytoncidal plants cleaning the air from microorganisms. Aspen (P. tremula L.) is the best known example of wild poplars or poplar trembling; it got its name from the constant thrill of leaves in the wind caused by the special structure of the stem, which is flattened in the plane. Genus Salix — Osier, it contains species sometimes easily interbreed, sometimes hardly distinguished. Male flowers of osiers have just 2–3–5, rarely up to 12, stamens, while the poplars have up to forty of them, at least 6 species of this genus are very diverse in form and leaf pubescence, on life forms. There are large shrubs such as osier ash (Salix cinerea), and tall, beautiful trees, such as osier silver (S. alba). Flowers of osiers are collected in inflorescence ament, lack of perianth, however pollinated by insects, as each flower, both male and female bears a honey piece of gland, which gives a rich nectar. Male flower of osier is in the axil of a silvery pubescent bract and consists often of 2, but in some species of 3–5 stamens, at the base of which lies a honey gland. Female flowers are also located in the axils of pubescent bract and consist of gynoecium with a clearly visible stigma and glands at its base. Silvery pubescent bracts create a silvery robe of familiar rams of osier. Fruit is represented by a capsule with numerous seeds and hairs. This genus is represented by goat willow (S. caprea), sharp-leaved willow (S. acutifolia). Babylonian willow with weeping branches (S. babylonica) is widely used for greening.

76 Order Nettles — Urticales. Trees, shrubs or grasses with alternate or opposite, simple leaves, usually with stipules, often accreted into caps belong to this order. The anatomy is characterized by the of lacticifers in some families or cystolytes in cells. The flowers are mostly in the inflorescences, sometimes have very complex structures, are only occasionally single, with a simple unprepossessing perianth, and sometimes it is completely lacking, 2–3–4-membered, usually unisexual, sometimes bisexual. The number of stamens is usually equal to the number of lobes of perianth, gynoecium consists of two carpels, ovary is inferior or superior. Nettles family (Urticaceae). Great nettle is a perennial dioecious herbaceous plant with long rhizomes. Stems are erect, dumb tetraquetrous, furrowed, with stiff stinging hairs. The leaves are opposite, ovate-lanceate, greatly digitate, covered with stinging hairs. Flowers are small, unisexual, green, with a simple perianth of 4 lobes. Male flowers have four stamens, female — a single pistil, sessile stigma. Inflorescences are axillary, long, spike-like, hanging. Fruit is represented by an egg-shaped nutlet. Its height is 30–150 cm. Dog nettle (burning nettle) is a monoecious annual grass with branched stems. The leaves are ovate-elliptic, sharp, incised-serrate, covered with stinging hairs. The flowers are small, green, with a simple perianth, stamen and pistil. The flowers are collected in spike-like inflorescence, which is equal in length to leaf petioles or shorter. For great nettle, in contrast to the dog nettle, spike-like inflorescences are hanging and longer than leaf petioles. Their height is 15–60 cm. One of the characteristic features of nettle is the abundance of cystolytes — whitish formations impregnated with calcium carbonate. Form of cystolytes (point, rod-shaped, oval, crescent, clavate, starry, F-shaped and etc.) is more or less constant for certain taxa, and often serves as a good diagnostic feature in the taxonomy of species and genera of the family. Burning nettle hairs have stinging cells (1 mg of its has nearly 100 stinging cells) containing corrosive liquid of complex chemical composition; it contains histamine, acetylcholine, formic acid. Burning hair looks like capillary tube terminating in a small round head. The upper part of the hair gets soaked with silicon and tt breaks off if touched, the sharp edges of the hair pierce the skin and inject the contents of stinging cells into the wound. The result is a painful burning — burn of a nettle. The representatives of this family are the great nettle (Urtica dioica L.) and burning nettle (Urtica urens L.). Elm family (Ulmaceae) — these are tree and shrub species with alternate or opposite leaves, are usually asymmetric and roughly pubescent. Trees are with a spreading crown and asymmetrical leaves — one half of the leaf is tapered at the bottom, and the second one is rounded and longer than the first.

77 The flowers are often collected in the small, inconspicuous inflorescence, unisexual or bisexual, 4–5-membered, with a free or slightly connate simple perianth, stamens are equal in number to the number of lobes of the perianth, sometimes adherent to it, gynoecium consists of two carpels, fruit is represented by a seeded nut sometimes winged or drupe. The representatives of this family are the different types of elm, which are found in their wild form, being a part of broad-leaved and spruce forests. European white elm (Ulmus laevis Pall.), naked elm (U. scabra). The flowers are small inconspicuous, have a well-marked perianth of 5–6 accreted leaves, 5 or 6 stamens, gynoecium is with large stigmas diverging in different directions. Even before falling off of the perianth but after pollination of flowers, on the sides of the ovary ribs appear, fast-growing, and in mature fruit they broad a wide wing surrounding the fruit. Ripening processes rapidly, and soon after the full deployment of leaves branches get covered with pale green, later yellowing fruits. These plants bloom until the full opening of leaves. Mulberry family (Moracae) — is the largest family in order Urticales, but in the temperate zone there is only a small number of species and those are mainly cultivated plants. These include woody plants and shrubs, evergreen and deciduous, the leaves are always with stipules, sometimes accreted into the cap. The anatomy characterized by cystolytes and lacticifers. The flowers are usually in inflorescences, small, inconspicuous, wind-pollinated and entomophilous, monoecious and dioecious. The shape of inflorescence varies: racemose, capitate, spicate, umbellate, inflorescence axis can be thickened, and take a spherical, saucer-shaped, the shape of a hollow sphere with flowers arranged in the inner cavity of the ball (seacon at Ficus sp.). Perianth consists of 2–8, more often of four leaves, free or slightly accreted, as many stamens as there are perinath lobes, or 3–2–1, gynoecium consists of 2 accreted carpels, the ovary is superior, semi-inferior and inferior. Fruits are drupes, in multiple fruits. The most famous is the mulberry (Morus sp.), or sycamine, of the family of mulberries bred in the southern regions of the country (in the Ukraine, Central Asia) for its sweet fruit is eaten in a raw or processed form, as well as for its leaves — food silkworm. There are commonly found two kinds of mulberry in a culture: white (Morus alba) and black one (M. nigra). These are tall trees with whole lobed leaves, flowering in the spring to complete their blooming. Male inflorescences of mulberry are amentaceous — like inflorescence of birches. On the elongated drooping axis, small greenish-yellow flowers with 4 tepals and four stamens are arranged on short stalks, opposing to tepals, in bud rolled inwards flower. Female inflorescences have much shorter axis sticking, not drooping, and here have tightly arranged small greenish flowers, bearing the same 4 tepals and gynoecium with a rather large globular ovary. Later, when the fruits ripen, tepals get fleshy, cover the fruit and get accreted to form

78 an elongated multiple fruit, slightly reminiscent a raspberry fruit. The plants are dioecious. One more interesting plant of this family is bread in the south — figs, or wine berry (Ficus carica L.), a tall tree with broad crown and large palmate- lobed leaves. The inflorescence of this genus is called seacon. The flowers are very small, white, with a poorly distinguishable perianth. Perianth has 5 leaves and 5 stamens and female flowers are of 2 types in male flowers sitting on marked pedicel: short styles and long styles, perianth has also 5 small leaves. Medicinal raw materials are leaves, fruits contain vitamins, enzymes. Other species of this genus — ficus (F. elastica) is widely known as indoor ornamental plants. Breadfruit (Artocarpus altilis), a major multiple fruit is a vital source of food for the residents of the tropical zone. Many mulberries, including mulberry (gen. Morus), gives fibers coming onto the ropes or paper, paper mulberry (Broussonetia papyrifera), some plants are used as as dyes the range of applications of the members is very wide. Hemp family (Cannabaceae) includes two genera: hemp (Cannabis sp.) and hops (Humulus sp.). These are herbaceous annuals and perennials. Genus Cannabis is represented by one species — common hemp (C. sativa L.), commonly known in the culture as a fiber, oil and narcotic plant. This is a fairly high dioecious erect plant with digitipartite leaves. Female separate instances of so-called fimble. They give a finer fiber, male — materka tougher. Male specimens of cannabis develop quite large paniculate inflorescences of yellowish-greenish quinary flowers. The inflorescences of female specimens are denser, each flower is enclosed in a folded coating sheet, spherical ovary carries two large stigmas, and a thin film — undivided perianth is worn in the bottom. Fruits are rich in oil, which previously was used as food and maintenance. The genus Humulus — contains only 2–3 species. It is well known as (H. lupulus L.) a perennial vine plant, with large palmate-lobed or digitipartite leaves, developing numerous glandular hairs, which are often used as an ornamental. The plant is dioecious. ―Cones‖, or rather the female inflorescences of hops are covered with capitate glands that release a resinous substance, the male flowers are very similar to the male flowers of cannabis. The flowers are inconspicuous, small, unisexual, wind-pollinated, collected in inflorescence. Male flowers have 5 tepals and 5 hanging stamens, female have split rudimentary perianth, gynoecium of two carpels, the fruit of a nutlet.

79 LECTURE № 9 A SYSTEMATIC REVIEW OF SUBCLASS ROSIDAE

Subclass Roses — Rosidae. Roses are the largest subclass of flowering plants that includes 40 orders, 160 families, almost 2800 genera and more than 55 000 species. They represent one of the main trunks of bipartite development, which is derived from the ancient magnolia, the most primitive representatives of Rosidae with actinomorphic flowers, which keep apocarpous gynoecium and an unindefinite number of stamens, resemble primitive groups of Dlleniidae. However, the majority of the Rosiidae has quite an affinity with Dilleniidae. Evolution of Rosiidae manifests itself in a gradual decrease of the number of flower parts, perianth accretion and formation of its zigomorfism, carpel accretion and formation of the inferior ovary with a single column. There are also examples of simplification of the flower and other plant organs due to the highly specialized, transition to anemophily or a dioecious type. The order of Saxifrage — Saxifragales. Includes 9 families. Family Jades (Crassulaceae) comprises about 35 genera and over 1500 species. These are mainly herbaceous, often partially woody leafy succulents with fleshy, sometimes cylindrical or spherical bluish leaves. Almost always bisexual, actinomorphic flowers of Crassulaceae are collected in terminal cymoidal inflorescences and usually brightly colored. Perianth is doubled, usually forms 4–5 free or intergrows lobes. Stamens are the same as lobes or twice as large, and their strands are free, or more often attached to the sides of flower tube. Gynoecium is apocarpous of several free or accreted at the very bottom of the carpel. *Са5Со5А5+5G5. Fruit is polyleaflet, rarely capsule, is formed with half –accreted leaflets. Crassulaceae are mostly monocarpous. Representatives — stonecrops species (Sedum sp.): common stonecrop (S. telephium L.), mossy stonecrop (S. acre L.), a large stonecrop (S. maximum (L.) Hoffm.), the grass of which contains flavonoids, phenols, coumarins and etc. Stonecrops are used in homeopathy. Broth or alcoholate tincture of Rhodiola rosea or golden root (Rhodiola rosea L.), has a bracing and tonic effect. Decorative representatives — houseleek (Sempervivum soboliferum L.), which form a dense rosette in the axils of leaves countless spherical ―children‖, easily broken off and rolled back to the side. Saxifrage family (Saxifragaceae) consists of 30 genera and about 600 species distributed in cold areas. All saxifrage — grasses, in a particularly severe climatic conditions forming dense cushion courtines. Bisexual or rarely unisexual flowers of saxifrages are single or collected in various cymoidal inflorescences. More often they are actinomorphic, rarely zygomorphic, when two neighboring lower petals are considerably larger than the others. Perianth is doubled, five-membered, but there are species whose petals may be completely reduced (golden saxifrage). Stamens are of 5–10, and

80 apocarpous or syncarpous gynoecium consists of 2–5 carpels. The ovary is superior, semi-inferior or inferior with a free, sometimes coalesce columns. *Са4-5Со4-5А5,8,10G(2-5)–. Fruit — capsule — opens by partitions. Representatives: marsh saxifrage (Saxifraga hirculus L.), golden saxifrage (Chrysosplenium alternifolium L.). Giant rockfoil (Bergenia crassifolia (L.) Fritsch), rhizomes contain up to 27 % tannins. Overwintered leaves serve as a substitute for tea, possess immunomodulatory effects. Family gooseberry (Grossulariaceae). The family comprises about 150 species. All members of this family are shrubs, flower structurally are very similar to the saxifrages. The flowers are usually collected in few-flowered axillary brush. Sepals of perianth accrete in the bases into a tube, and the petals are strongly reduced. Stamens are alternate with petals. Syncarpous gynoecium consists of 2 carpels; it forms the inferior ovary which develops into a juicy berry. Polymorphic genus currant (Ribes sp.) is interesting, some species of which are sometimes isolated in a separate genus gooseberries (Grossularia sp.). All known kinds of currant are derived from black currant (R. nigrum l.). Red currant (R. rubrum L.), pigweed (R. sativum Syme) are widely spread. Common gooseberry (Grossularia reclinata (L.) Mill.) differs from the prickly currant with pubescent berries and prickly shoots. The fruit of a black currant is used in multivitamin fees, the leaves are used as a spice. The order Roses — Rosales. Unite trees and shrubs, deciduous and occasionally evergreen plants, as well as numerous herbaceous plants. Leaves are simple, alternate or opposite, variously dissected, complex and secondarily simple, usually with stipules. Segments of vessels have a ladder and a simple perforation. Flowers are bisexual, sometimes unisexual, mostly actinomorphic. Androecium varies from numerous and uncertain to numerous and definite. Gynoecium is apocarpous, syncarpous and monokarpous, the ovary is superior or inferior. Fruits are different. The order includes three families, of which the largest by number of delegates (which also has a great practical importance) is the family Rosaceae — roses, widespread in the temperate zone, and represented in the tropical and subtropical zones. Family Roses (Rosaceae) includes about 100 genera and over 3000 species, widespread everywhere. This family is represented by trees, shrubs and herbaceous plants. Flowers are actinomorphic, bisexual with a double 5-membered perianth. Androecium is numerous, number of stamens is is two- four times higher than than that of petals. For the flowers of the whole family there is a typical part — hypanthium — extended receptacle, the bases of the stamens, petals, sepals adhere to the edges of it. Often hypanthium has a cup-shaped form that facilitated the development of the bottom ovary in this family. Gynoecium and fruits of the Rosaceae vary, that is why the family is divided into 4 subfamilies.

81 Subfamily Spiraeoideae is characterized by a flat hypanthium and apocarpous gynoecium. Fruit — polyleaflet. In the temperate zone this subfamily is represented by shrubs of genus spiraea (Spirea sp.): willow spiraea (S. salicifolia L.), oriental spiraea (S. media L.); genus false spiraea (Sorbaria sp.), physocarpus (Physocarpus sp.). Subfamily Rosa (Rosoideae) is characterized by a convex hypanthium, apocarpous gynoecium. Fruit — polynutlet. Separate fruitlets are located on the lush (strawberries), or at least within a juicy (rose hips) receptacle. In addition to these kinds of fruits are juicy drupes (raspberries, blackberries). Representatives: different species of bloodroot (Potentilla sp.), cuffs (Alchemilla sp.), bennet (Geum sp.), agrimony (Agrimónia eupatória), raspberry common (Rubus idaeus L.). Leaves are imparipinnate quinary, fruit is polydrupe; roebuck berry (Rubus saxatilis L.), weak creeping stems, trifoliate leaves, fruit is polydrupe; blackberry (Rubus caesius L.), leaves are tricompound, petals are running a part; strawberry (Fragaria vesca L.), creeping stems, trifoliate leaves in the socket, the fruits — polunutlets; garden strawberry (F. ananassa); cultural strawberry (F. moschata Duch.); marsh cinquefoil (Comarum palustre L.), arcuate rhizome, odd-pinnate leaves, by the structure of flowers and fruits it is similar to wild strawberry. The plant is poisonous, is used for treating cancer goose foot or silverweed cinquefoil (Potentilla anserina L.), stem is creeping, unpaired — interruptedly pinnate leaves, flowers are 5-membered; erect cinquefoil (P. erecta (L.) Rausch.), the stem is erect, the flowers are 4-membered; evans’-root (Geum rivale L.), leaves are intermittently lyrate. There is a ginophore; meadowsweet (Filipendula ulmaria (L.) Maxim.), leaves on the bottom have dense white hairs, inflorescence — antella. Subfamily plum (Prunoideae) is characterized by cup-shaped hypanthium, monokarpous gynoecium. Fruit — juicy drupe (plum, cherry, peach, apricot, cherries, etc.) or dry drupe (almonds). Most of the plums — plum, cherry, cherries, apricots and others have a solid stony endocarp, which is surrounded by rich, brightly colored mesocarp. The tree and shrub species with simple leaves, stipules, usually falling down early belong to that subfamily. The flowers have five fingers of the structure plan, with a simple cup, numerous androecium and monomeric gynoecium, sitting at the bottom of cup-shaped hypanthium. Most members are crop plants, also include many drugs. Representatives: bird cherry (Padus avium (P. racemosa (Lam.) Gilib.), the fruits of which contain tannins. Sour cherry (Cerasus vulgaris Mill.). Subfamily apple (Maloideae) — characterized by the fruit of an apple or little apple produced by fusion of the walls of the hypanthium’ carpels (inferior ovary), which often grow up and become juicy. This subfamily is represented by cultural and wild species: cultivated apple (Malus domestica Borkh.), crab apple (M. silvestris Mill.), choke pear (Pyrus silvestris L.), field ash (Sorbus

82 aucuparia L.) the fruits of which contain many biologically active substances used in homeopathy, bloody-red hawthorn (Crataegus sanguinea Pall.), flowers and fruits of which are used for diseases of the cardiovascular system. Family oleaster (Elaeagnaceae) — low trees or shrubs, whose shoots and leaves are covered with thyroid or star, silver or brown scales. Leaves are alternate or opposite, without stipules, whole and entire. Flowers are bisexual or unisexual, regular, with a simple perianth tube, positioned by one or by few- flowered corymbose bundles in the leaf axils. Perianth is 4-lobed, sometimes with a different number of leaves. There are four-twelve staments. The ovary is superior with hypogynous disc that grows up in fruit, 2-alopecia. Fruit is juicy or dry, kind of drupe, with 1 seed. A characteristic feature of the oleaster is the presence of tubercles on their roots with nitrogen-fixing bacteria. Due to the presence of bacteria, these plants are able to improve even very poor soil. Representatives: sea buckthorn (Hippophaë rhamnoides) is used to produce oil, which has a regenerating effect and is used for the treatment of venous ulcers and burns. Order Legumes (Fabales) includes three families: Mimosa, Caesalpinaceae and actually Legumes. Representatives of this order are woody plants, shrubs, vines and numerous grasses, with leaves mostly complex with stipules. The order is distributed very widely, from the Arctic to the tropical zone and is of considerable practical importance, since it has a lot of useful plants: food and fodder plants that accumulate the protein, numerous ornamental, medicinal, spicy, poisonous, giving gum and valuable wood. The families included in the order differ by the structure the structure of the flower, a 4-5-membered flower with numerous, sometimes reduced to 4 stamens, androecium is typical for the most primitive family of Mimosa. In other families androecium is already established and is in most cases 10-membered. The number of carpels varies from 5 to 1, the fruit is a bean or beans. Family — Mimosa (Mimosaceae). In order Fabales this family is the most primitive. Its members are mainly woody plants (sometimes water (Neptunia sp.) With decompound or twice decompound leaves, which when touched can fold (shy mimosa), with stipules, that for some species are transformed into spines. Sometimes the leaf is reduced and replaced with phyllodes — advanced petiole. Flowers are 4–5-membered, always actinomorphic, with a fine doubled perianth, sometimes accreted at the base, usually not large, clustered in spherical or more elongated, resembling brush, inflorescences. Inflorescences are made notable due to the numerous, long, painted in yellow, pink, lilac, purple color stamens, sometimes free, sometimes accreted. Gynoecium in the most primitive forms is of 5 or 2 carpels giving after maturation 2–5 beans, and the majority has of one carpel gynoecium and fruit bean, while some mimosa can have a giant one — up to 1.5 m (Entada sp.).

83 These plants are characterized by a selection of gums, such as plum gums that are used in the industry (gum arabic). The most familiar are two species of ―mimosa‖ (Acacia dealbata L.), black locust (Robinia pseudoacacia L.) contain essential oil in its flowers. Family Caesalpiniaceae. To this family belong mainly woody plants, shrubs, vines, rarely herbaceous plants. Leaves are compound, rarely simple, with stipules. The flowers within the family show a gradual transition from actinomorfism to zygomorfism. Perianth is double, 5-membered, can be reduced. Androecium has 10 stamens lying in the two circles, but often part of the stamens is transformed into staminodes; gynoecium has one carpel, giving the fruit of the bean. Beans of Caesalpiniaceae are often large, cylindrical or flat, sometimes fleshy. Representatives: Indian senna (Cassia acutifoilia L.), the leaves of which are part of antihemorroidal and laxative fees. Family Legumes (Fabaceae, Leguminosae, Papilionaceae). Tree, shrubs, vines and numerous grass plants belong to this family. Herbaceous forms are typical for countries with a temperate climate. The leaves of beans are complex: pinnate, palmate, trifoliate. The flowers are often of five-typed plan with double perianth, always zygomorphic and mostly with a butterfly whisk, whose upper lobe — sail is larger than the others, two petals — oars, and two accreted petals form a boat. Androecium is of 10stamens, 9 stamens are accreted into a tube and only one remains free ↑Ca5Co2+2+1А(9)+1G(1). Sometimes all 10 stamens are accreted. The gynoecium is of 1 carpel, always giving after ripening a fruit bean. Most beans are opened with two doors, scattering seeds; for clovers number of ovules and seeds is cut and the fruit stops disclosing. There are a lot of modifications of beans in this family. Representatives: king’s-clover (Melilotus officinalis (L.) Desr.), field pea (Pisum sativum L.), Russian beans (Faba vulgaris Moench), yellow acacia (Caragana arborescens Lam.), kidney bean (Phaseolus vulgaris), craw pea (Lathyrus pratensis L.), red clover (Trifolium pratense L.), purple alfalfa (Medicago sativa L.), common licorice (Glycyrrhiza glabra L.) etc. Legumes are of great practical importance. The presence of nitrogen-fixing nodule bacteria on the roots enhances soil fertility, protein accumulation in seeds. Plants of this family are widely used as green manure, for decorative purposes, as medicinal plants. The practical value of legumes is huge: large and bright colors make them desirable ornamental plants. Medicinal plants: Glycyrrhiza glabra and G. Uralensis, Thermopsis, Melilotus officinalis Cultivated plants include a pea Pisum sativum. It has pair-decompound leaves (part of them is turned into tendrils with leafy stipules); dyeing: Indigofera tinctoria, Genista tenctoria, giving dye indigo and basma. Russian beans Faba vulgaris is an annual plant with ternate leaves, flowers are in axillary racemes, beans are large, hairy.

84 Beans Phaseolus vulgaris. Leaves are trifoliate, yellowish-white flowers, the usual structure of the legume. Seeds and unripe beans are used in the food. Soybean Glycine max, trifoliate leaves, yellowish flowers. Ornamental plants: sweet pea Lathyrus odoratis, lupine Lupinus polyphullus with palmately compound leaves and androecium having only 10 stamens, they contain toxic alkaloids. Yellow acacia Caragana arborescens — tall shrub with unpaired-pinnate leaves and stipules of thorns. Lathyrus pratensis, has a pair of leaves with a mustache, one-sided brush. Vic — Vicia cracca and V. sepium. V. cracca, or mouse peas, with numerous tendrils of many pairs of leaflets, leaves, one-sided brush. Trifolium pratense, red clover, T. repens, white clover, and T. Hybridum, hybrid clover, all clearly visible stipules, inflorescences in the form of the capitulum, the fruits - small, indehiscent legumes. Alfalfa Medicago sativa, is a perennial plant with ternate leaves and brush blue-violet flowers, also a medicinal plant. M. officinalis, medical melilot. Ginseng — Panax ginseng — a medical plant with palmate-lobed leaves, a fleshy root. Eleuterococcus senticosus — a large spiny shrub with high palmate-lobed leaves, inflorescence is an umbrella, berry-like fruit. The extracts have toning properties. Order Araliales includes 2 families. Life forms are represented by woody plants, shrubs, vines and grasses, often with alternate, rarely opposite leaves with or without stipules. Flowers are 5–4-membered, bisexual, with a double perianth, but with a much reduced calyx. 5–4 stamens, gynoecium is of two carpels, ovary is inferior Ca4-5Co4-5А4-5G(2). The small flowers are collected in umbellate inflorescences. Fruits are berry-like or drupes, Umbrellas have cremocarp. Family Aralia (Araliaceae) — woody plants, shrubs, vines and perennial herbaceous plants with alternate, entire, simple, one-piece, palmate-lobed or digitipartite leaves with small stipules. The stems have secretory channels. The flowers are collected in an inflorescence umbrella or capitulum. Flowers are 5-membered, with little noticeable calyx and 5–10 petals. 5 stamens, gynoecium is of 2–5 carpels, the ovary is inferior with glandular disc. Ca5Co5-10А5G(2-5). Fruit is represented by an apocarpous berry or drupe. Representatives: ginseng (Panax ginseng C. A. Mey), eleuterococcus (Eleutherococcus senticosus L.), Manchurian aralia (Aralia mandshurica L.) (the roots of which are used as a tonic and adaptogenic remedia). Family Umbellate (Apiaceae or Umbelliferae) — herbaceous plants, annuals and perennials. Umbellate are large plants with a powerful and ribbed hollow stem, large highly dissected leaves with large sheaths concealing inflorescence. Inflorescence is simple or complex, either umbrella or capitulum. 85 Flowers are actinomorphic or zygomorphic at the edges of inflorescences, with almost imperceptible cup of 6 teeth, 5 petals and stamens and gynoecium of two carpels, and an inferior ovary bearing nectariferous disc. (Ca6Сo5А5G(2)). Fruit is represented by a cremocarp, rarely a drupe. Fruitlets bear 5 main ribs on its surface, between which there are oil channels. In the tissues there are secretory canals. Umbellate are traditional crops. Carrot (Daucus carota L.) — biennial plant. Its blossoms have very large feather wrap, that distinguish this genus well from other umbellate. The fruits are used in folk medicine for the expansion of the coronary vessels, daucarin is prepared. Dill (Anethum graveolens L.), the leaves are cut into narrow thread-like lobes. Caraway (Carum carvi L.) has twice — thrice pinnate leaves, relatively small, it is characterized by pinkish flowers and fruits of aromaticity with a characteristic odor. It is used in baking and as a medicinal plant, its fruits have anti-inflammatory action. Cowbane (Cicuta virosa L.), which is characterized by tuberous rhizome, the cut having a cell filled with a poisonous juice, smooth stem, leaves tripinnate. Hemlock spotted or spotted omega (Conium maculatum L.), has a mouse smell and purple spots or streaks on the stem. Trifoliate leaves, the flowers are white. Lovage (Levisticum officinale), has a specific smell, which is a bit like the smell of celery. Hence, one of the popular names — winter celery. Lovage stimulates the appetite, relieves intestinal cramps, has a carminative and diuretic effect. Order Dipsacales includes 8 families and is represented by woody plants, shrubs with a whorled, entire, lobed, simple leaves without stipules. Flowers are zygomorphic, 4–5-membered, hamopetalous. Androecium is equal in number with patals or less, gynoecium has carpels, the ovary is inferior, ↑Ca(4-5)Co(4-5)А4-5G(2), fruit is represented by a capsule, drupe, berry or nutlet. Honeysuckle family (Caprifoliaceae) includes 13 genera and 300 species, represented by shrubs or small trees, sometimes shrubs, grasses or shrubs with opposite, simple or complex leaves without stipules. Flowers are 4–5-membered, hamopetalous. A cup is medium-sized, cluttering the inferior ovary, sometimes two-lipped corolla, campanulate or rotate. Androecium has 3–5 stamens, adherent to the corolla, gynoecium is of 2–5, rarely 8 carpels with uneven development of nests of the ovary, the ovary is inferior, sometimes semi-inferior (↑Ca(4-5)Co(4-5)А3-5G(2-5)), flowers are collected in inflorescence of cymose. The fruits are represented by berries, drupes, capsules. Representatives: common elderberry (red) (Sambucus racemosa L.), black elderberry (Sambucus nigra), cranberry tree (Viburnum opulus L.), the fruits of which contain cyanogenic glycosides and are used as a laxative and an emetic; common honeysuckle (Lonicera xylosteum L.), fruits contain essential oils and irridoidy has anti-spasmodic action; Linnaeus northern (Linnaea borealis L.), a small, creeping shrub in the moss.

86 Valerian Family (Valerianaceae) includes 13 species and 400 genera. Mainly grasses, sometimes shrubs and shrubs with opposite, whole or pinnate leaves belong to this family. Flowers are bisexual or unisexual, sometimes actinomorphic, but in most cases asymmetrical, 5-membered, calyx has 5 blades or jags, but sometimes by the flowers in the form of a ring, and in the fruit it is growing and serving for distribution, corolla is tubular, often with a medium- sized spur or of the bag-shaped process, emphasizing the asymmetry of the flower. Androecium is of 4–3–1 stamens, adherent to the corolla, gynoecium is of three carpels, but develops only one of them, the ovary is inferior (Ca(5)Co(5)А4-3-1G(3)). The small flowers are collected usually in polyflowered, complex inflorescences. Fruit is dry, nutlet-kind often provided with pappus formed of a cup. The most important from a practical point of view and the best known species of this family is a medical valerian, (Valeriana officinalis L.), whose strongly smelling rhizome is well-known for the drug.

87 LECTURE № 10 SYSTEMATIC REVIEW OF THE SUBCLASSES LAMIIDAE, ASTERIDAE

The Subclass Labiatae — Lamiidae. Labiatae originated from the representatives of the subclass Rosidae and represents the strong evolutionary branch, which characterized by high specialization of gamopetalous, usually tubular perianth. They contain 11 orders, 51 families, about 2400 genera and almost 40 000 species. This subclass is represented by trees, shrubs and herbaceous forms. The certain number of members of perianth, the strengthening of zigomorfiya on the way to specialization to the cross entomophilous pollination are typical for the flower of all species labiatae. The syncarpous gynoecium consists of 2 the carpels, that form superior, half-inferior ovary or inferior ovary. The order Gentianales. This order includes 13 families. All local representatives of this order are heliophilous herbs with opposites, entire leaves and with rather large brightly colored flowers in thyrsoid inflorescences of a different shape. The flowers are usually bisexual, actinomorphic or little zygomorphic, almost always 4–5-membered, with half-fused sepals. The family Rubiaceae is the most large family in this order. The family Rubiaceae includes about 500 genera and almost 7000 species, that are found all around the world. Rubiaceae are represented in the form of trees, shrubs and woody vines in tropical zone and in the form of perennial and annual grasses in the temperate zone. The opposite or whorled leaves with stipules are often fused, in the form of cap and then may fall off or stay in the form of trumpet. The flowers are big, single or small, in botryoid, paniculata or spherical inflorescence, where the flowers may stay separately from each other or fuse with each other. They are usually bisexual, rarely uniseaxual, actinomorphic or little zygomorphic, with underdeveloped, more often 4–5-toothed brightly colored calyx and gamopetalous 4–5-lobed, usually trumpet corolla. The inflorescence sometimes surrounded by the big colored bracts. The stamens attached to the tube of corolla and alternate with the petals. The syncarpous gynoecium consists of the 2 carpels, that form lower and very rare half-inferior ovary, 1–2-locular ovary on the top with nectar disk and long column, that is carried multibladed stigma.*Са0Со(4)А4G(2). The fruits are brightly colored berries or drupes, that are formed the stem or capsule and also dry fractional fruits. The seeds and dried fruits often have the pappus or clinging spines. The most widely distributed plants are representatives of the order Coffee (Coffea sp.): Coffea arabica (С. arabica L.), 2 species of Cinchina: Cinchona ledgeriana Moens ex Trimen, C. succirubra Pavon, their cortex contains the alkaloid quinine, Carapichea ipecacuanha (Psychotria ipecacuanha L.), and

88 also the representatives of the genus Galium (Galium sp.): Northern bedstraw (G. boreale L.), sweet-scented (G. odoratum L.), common marsh-bedstraw (G. palustre L.), yellow bedstraw (lady’s bedstraw) (G. verum L.), fen (G. uliginosum L.), Dyer’s madder(common madder) (Rubia tinctórum L.). The family Gentianaceae includes 83 genera and more than 1000 species. The representative of this family widely distributed everywhere but mostly in the temperate and subtropical zones. Gentianaceae represented by heliophilous perennial and annual grasses with the opposites, entire leaves. The flowers are big, brightly colored, bisexual, actinomorphic or little zygomorphic, 4–5- membered. The corolla’s petals are half-fused or just fused (tubular corolla with the twisted in the bud limb). The stamens alternate with the petals and attached to the its tube by thin threads. The syncarpous gynoecium consists of 2 the carpels, that form superior unilocular ovary with the sessile stigma or with the column, that is carried two-bladed or capitate stigma. The flowers have nectaries and nectar glands in form of fringed pocket, that are situated in the corolla’s tube *Са(4-5)Со(4-5)А(5)G(2). The flowers may be in thyrses of a different shape. The fruits are expanding polyspermous capsules, rarely berrylike. The seeds with the endosperm and little ovule, sometimes with not big pappus. Among Gentianaceae such species are notable: the gentiana (Gentiana sp.): the cross gentian (G. cruciata L.) and the marsh gentian (G. pneumonanthe L.). Their radices contain bitterness and are widely used in homeopathy. The Swertia perennis (Swertia perennis L.) is often found in the damp places The annual representatives of the genus Centaury (Centraurium sp.), that are used in medicine, can be found in the dry meadow. There are a lot of officinalis, beautifully blossoming, ornamental plants among Gentianaceae, that are used in the rocky gardens and in the alpine gardens. The family Menyanthaceae includes 5 genera and 50 species, closely adjoins to the Gentianaceae. The representatives from the family Menyanthaceae are water or coastal-water plants with opposite, simple, without stipules leaves. The flowers are big, brightly colored, regular, bisexual, 5-membered. Sepals are accreted at the base, petals form short tube. The valvate folded petals are in the flower’s bud. The stamens (5) accreted with the corolla’s tube. There are nectaries. The syncarpous gynoecium consists of the 2 carpels. They form superior 1 locular ovary. The fruit is syncarpous capsule. The most famous representative of this family in the local flora is Menyanthes trifoliatа L. It is a coastal or marsh perennial plant with a rhizome. The leaves are simple, ternately-dissected, macropodous, regular and sheathing . Ca(5)Co(5)А5G(2), the flowers are in raceme. The fruit is a capsule. The Menyanthes trifoliatа plays a big role in lots of types of bog communities and in the overgrown ponds, participating in the formation of quagmire. The leaves are the crude drug. They are used in form of infusion, gathering as the stimulating the appetite, cholagogue remedy.

89 The family Apocynaceae includes about 300 genera and more than 1500 species, most of them are woody vines, sometimes the trees, shrubs and herbs can be found. The leaves are entire, opposite, rarely in whorls or regular. The availability of the latex is typical for the all organs of the plant. It is usually has a rubber. The flowers are bisexual, actinomorphic, 5-membered(very rare 4-membered). They are in cyme inflorescence. They are rarely situated singly on the shoot tips or in the leaf axils. The calyx is usually cut almost to the base and a corolla often is tubular, rarely is patellate with the part of bending, rolled in the bud. Squamose or petaloideous appendages, that are shown from the pharynx, often located on the inner side of corolla’s tube. The stamens alternate with the petals and attached to the its tube by thin threads. The syncarpous gynoecium consists of 2 the carpels(very rarely of 3–5), sometimes can be secondary apocarpous. The fruits in most cases are consists of 2 follicles, that are fused on the base, opened at the seams. The seeds have the pappus from the silk hair or from filmy fringe. There are juicy indehiscent fruits that are spreaded by animals. In some cases the fruit can be capsule-shaped or indehiscent at full intergrown of follicles entired. Sometimes, the fruit is covered by the prickles or by the clinging outgrowths. The representatives are: the Strophanthus gratus (Hook) Franch, that contains cardiotonic glycosides; the Indian snakeroot (sarpagandha) (Rauwolfia serpentina Benth.), their radices contain alkaloids; periwinkles (Vinca sp.): the large periwinkle (V. major L.), the dwarf periwinkle (V. minor L.), the rosy periwinkle (Catharanthus roseus (L.) G. Don f. = V. rosea (L.)), that contain alkaloids. The order Oleales. The order includes only 1 family. The family Oleaceae includes 30 genera and about 600 species, that can be found all around the world. All representatives are woody vines-shaped or erected shrubs or trees with opposite (rarely regular), simple, trifoliate or odd pinnate leaves. The flowers are big, actinomorphic, brightly colored, bisexual (rarely unisexual), gathered in the racemosa or paniculata inflorescences. The calyx is absent or consists of 4 jags, that are fused at the base. The corolla is usually large or funnel-shaped with 4 parts, rarely dissected almost to the base. In some cases corolla and calyx are reduced (ash-tree) because of the transition to the anemophily. In this case, stamens departs from the receptacle and in case of corolla’s availability, they attach to the tube’s walls. There are 2 stamens (rarely 3–5). The syncarpous gynoecium consists of 2 the carpels, that form superior 2 locular ovary with bilobed, sometimes sessile stigma. Ca(4)Co(4)А2G(2). The fruits are transversely dehiscing capsules, samaras, berries or drupe, that are often monospermous. The seeds in the capsules often have the pappus and spreaded by the wind.

90 The representatives are: the European ash (Fraxinus excelsior L.), the Manchurian Ash (F. mandshurica L.), the mountain Ash (F. rhynchophylla L.), the South European flowering ash (the manna ash) (F. ornus L.), its cortex is used in case of diarrhea, dysentery as the anthelmintic, tonic and astringent remedy. Olea sylvestris (Olea europaea L.), its fruits are crude drug, that are used in producing the edible and medical oil. The order Solanales. The order includes 5 families. The family Solanaceae the biggest family in this order, includes 90 genera and almost 3000 species. The representatives are herbs, shrubs or little trees, sometimes epiphytic and parasitic vines. The leaves are opposite entire or odd pinnate. The flowers are bisexual, actinomorphic or little zygomorphic, gathered in cymose inflorescences. They have 5-lobed calyx and sympetalous rotate or tubular, 5-jagged corolla, that as an exception may be a little bilabiate. There are 5 stamens (rarely 2–4). They are often snugly to column and form the similarity of the cone. The syncarpous gynoecium consists of 2 the carpels(rarely of 5), that form superior 2-locular ovary with simple column and 2-lobed stigma. Ca(5)Co(5)А5G(2). The fruit is a berry or a dehiscent capsule. The representatives are: the bittersweet nightshade (Solanum dulcamara L.), that is used in homeopathy, the black nightshade (S. nigrum L.), the fruits is used as the antiseptic, diuretic and laxative remedy. The Deadly Nightshade (Atropa belladonna) is widely used in medicine as the spasmolytic remedy. The Jimson weed (Datura stramonium L.), the black henbane (Hyoscyamus niger L.). All parts of this plants contain alkaloids, the leaves are used in anti- asthma gatherings. The strawberry groundcherry (Chinese lantern) (Physalis alkekengi L.), is cultivated as ornamental, edible and officinal plant. The fruits contain a significant amount of ascorbic acid and the seeds contain up to 30 % fatty oil. The potato (Solanum tuberosum L.), the tomato (Lycopersicon esculentum L.), Capsicum annuum L., the Eggplant (Solanum melongena L.), are of great value as the food, containing vitamins, microelements and other substances. The Tobacco plants (Nicotiana tabacum L.) contains poisonous alkaloid, that is called nicotine, in its leaves. The Aztec tobacco (N. rustica L.), its leaves contain citric acid and nicotine, but the amount of the nicotine is much lower than in tobacco’s leaves. The order Convolvulales. The order includes 2 closely related families. The family Convolvulaceae includes 58 genera and about 1700 species, that are widespread. They are annual and perennial, decumbent and curling, sometimes dwarf shrubs, shrubs and even small trees with opposite, superior, pinnate or digitate, often with tomentous leaves. The flowers are brightly colored, bisexual (sometimes unisexual-in that case the plant is dioecious), actinomorphic, 5-membered. They are gathered in axillary inflorescences, apical, rarely the flowers are single, axillary. The sepals

91 are free, some of them are greatly grown. The corolla is sympetalous, infundibuliform or tubular. The stamens alternates with petals, that are attached to the base of the corolla’s tube. The syncarpous gynoecium consists of the 2 carpels (rarely of 3–5), that form superior 1–2(3–5)-locular ovary with 1–2 with ovules in each locule. Columns almost free or fused to form a common 2-lobed or capitate stigma. Ca(5)Co(5)А5G(2). The fruit is a capsule, indehiscent or juicy in the form of nut. Convolvulaceae seeds are covered with hairs that facilitates the transfer of the wind. Many species convolvulaceae are decorative and they are often grown in greenhouses. The field bindweed (Convulvulus arvensis L.) is the climbing plant with quite large pink and white flowers, that is used in homeopathy. Ipomoea purga (Wend.) Hayne — roots contain glycosides resin that is used as a purgative. Sweet potatoes or yams (Ipomoea batatas (L.) Lam.), starch from thickened lateral radices, known as the ―Brazilian arrowroot‖, is used as enveloping and emollient remedy. The order Boraginalеs. There are 7 families in this order, the largest of which — the family Boraginaceae. The family Boraginaceae includes about 100 genera and more than 200 species that are found almost everywhere. Predominant among them the grass, but there are also shrubs, bushes and even small trees, annual and perennial grasses. Leaves are entire, alternate have bristly pubescence. Flowers actinomorphic or slightly zygomorphic, usually bisexual, five-membered. The calyx is sympetalous, can grow strongly in fruits. The corolla is also sympetalous rotate or roll. Stamens alternate with corolla’s lobes and are attached to its pipe, sometimes fused, surrounding the column. The syncarpous gynoecium consists of 2 carpels, forming the superior 2 locular ovary, which nests are often separated the false wall in such way, that the ovary is divided into 4 Secondary camera with a single ovule in each nest. The column with two-bladed or capitate stigma, coming out of the recesses between the blades of the ovary, fused only at the bottom. The flowers are collected in monochasial cyme, that can form more complex paniculate, corymbose, spike and even capitate inflorescences. The fruit is dry, fractional, decays into 2–4 indehiscent share called Eremo. Seeds and fruits are often covered with growths clinging, sometimes equipped with fleshy appendages that attract ants. The representatives are: forget-me-nots (Myosotis sp.) —M. palustris Lam., M. arvensis (L.) Hill, the Lungwort (Pulmonaria obscura Dum.), the wood-mat (Cynoglossum officinale L.), the knitback (Symphytum officinale L.), its radices contain alkaloids. The order Scrophulariales. There are 15 families. We will review two families (Scrophulariaceae and Plantaginaceae) that can be found in Belarus. The family Scrophulariaceae is in the center of the order. There are about 300 genera and more than 5000 species of plants. Annual grasses dominate in

92 the family, and small shrubs and shrubs are found. Leaves are alternate, rarely opposite or whorled, without stipules. Flowers are bisexual, zygomorphic, collected in terminal and axillary inflorescences, or located on one in the leaf axils. Corolla and calyx consists of 4–5 lobes, that are often intergrow into bilabiate tube, often having saccular bulge or spurlike. Flowers with two-humped coolla adapted to pollination by large insects, which can bend its weight lower lip and access the nectar, which is located at the bottom of the corolla’s tube. There are 4 or 2 (rarely 5) stamens in the flower. In case of developing 4 stamens, there is often 1 major staminode. Stamens are usually arranged in two pairs of unequal development and are attached to the corolla’s tube. One pair of stamens falls and remain in bloom for only 2 stamens in some figwort, for example, Veronica. Syncarpous gynoecium of 2, rarely 3–4 fused carpels forming the superior 2 locular ovary with whole or 2-lobed stigma. Fruits are capsules, they are very rare berries — or stone-like. The seeds are often surrounded by membranous border and spreaded by the wind. They can be taken away by the ants, attracted by the school oily appendage The representatives are: the variety of foxgloves (Digitalis sp.): the purple foxglove (D. purpurea L.), the Big-flowered foxglove (D. grandiflora Mill.), their leaves contain cardiotonic glycosides. The eyebright (Euphrasia officinalis L.), its herba is used as трава которой используется как anti-catarrhal and astringent remedy. Melampyrum nemorosum L., Pedicularis palustris L., the woodland figwort (Scrophularia nodosa L.), the great mullein (Verbascum thapsus L.), their leaves are used as the expectorant and emollient remedy in diseases of the superior respiratory tract. Veronica chamaedrys L., the common toadflax (Linaria vulgaris Mill.). The family Plantaginaceae — widely spread, included 3 genera and about 300 species, that is familiar to Scrophulariaceae. Mostly, they are small rosette grass, although there are sometimes small shrubs. They are characterized by capitate or spike inflorescences with small wind-pollinated actinomorphic bisexual flowers having 4 sepals, 4 petals and 4 stamens. Syncarpous gynoecium of 2 carpels form the superior 1–2 locular ovary, that is developed in capsule, that is opened by round lid. Ca4Co(4)А4G(2). The typical representer is the broadleaf plantain (Plantago major L.) and related species are used as antibacterial, wound healing and expectorant remedy. The order Lamiales. This small order close to Scrophulariales, but more organized and its morphological characteristics are more determined. In the main, there are perennial herbs, shrubs or bushes, sometimes woody plants with simple opposite leaves without stipules. Flowers are always zygomorphic with the fused calyx and a corolla, the androecium of 2–4 stamens, sometimes 5, gynoecium of 2 carpels, the ovary is superior, the fruit is stone-like or fractional 4-nut.

93 There are 3 families. The family Lamiaceae or Labiatae is one of the largest families of flowering plants. To this belong about 200 genera and over 3500 species, that are widely spreaded in all climatic zones. The representatives sometimes form cushion strongly spiny clumps in desert areas. Herbaceous plants dominate in the family, although there are bushes, shrubs, sometimes trees with simple, sometimes complex leaves and often quadrangular stem. The flower’s corolla is tubular, two-lipped, is formed by, almost always five, rarely four colored petals. The superior lip is formed by the 2, and the lower by the 3 petals, although there may be exceptions. For example, the mint flowers are actinomorphic. Calyx, remaining in fruit, is formed by the 5 fused sepals. It can be painted, but often the calyx is green, and its various modifications are associated with with the adaptation to spread by the wind or animals. Calyx jags can turn into clinging hooks or bent blade, increasing windage. There are usually 4 stamens in two unequal pairs in the flower, one of which can be strongly reduced or even completely reduced. Stamens often have a rather complicated structure, which is determined by the adaptation to pollination by certain types of insects. At Sage, for example, stamen resembles the lever, providing a rash on his back insect pollen, sticks his head in the corolla tube in search of nectar. Syncarpous gynoecium consists of two carpels forming the top 2-well ovary. Speaking about sage, for example, stamen resembles the lever, providing the rash of pollen on the insect’s back, sticks its head in the corolla’s tube in search of nectar. Syncarpous gynoecium consists of 2 carpels, that form the superior 2-locular ovary. Both nests are separated by a false wall socket to 2 chambers, each of which contains one ovule and that is why the gynoecium is 4-bladed. The coenocarpous fruit decays into 4 mericarps (Eremo) nut. Separate semi mericarps often have their own adaptations for dissemination — hairs or crests or spread with a calyx in which they are connected by a ring of hairs on an inner surface of its tube for a some time. Flowers are bisexual, collected in complex inflorescences with shortened secondary axes, that is why they look spiky. Quite often the main axis of the inflorescence is greatly shortened and then, and then it takes the form of the head. All parts of the plants contain various essential oils. There are a lot of many valuable representatives among the plants of this genus: Lavandula sp., Rosmarinus sp., the mint (Mentha sp.), the balm mint (Melissa sp.), the spurflowers (Plectrantus sp.) and Pogostemon sp. The oils, obtained from them, have antibacterial properties and widely used in medicine, cosmetics and perfumes. Some species, especially the noble basil (Ocimum basilicum L.), are cultivated as food, spice plants. Young shoots of the representatives of the genera mint, thyme are used as fragrant spices (Thymus sp.), the savory (Satureja sp.), and marjoram (Origanum majorana L.). Chinese artichoke has edible tubers(introduced into the culture in China) (Stachus affinis L.).

94 As a medicinal plant raw materials in the GF RB are indicated: lavender flowers (Lavandula officinale), oregano’s herb (Origanum vulgare), sage leaves (Salvia officinales), peppermint leaves (Mentha piperita), herb of the lemon balm (Melissa officinalis), indian kidney tea leaves (Ortosiphon stamineus), thyme herb (Thymus serpyllum) herb thyme (Thymus vulgaris). The subclass Asteridae. The subclass Asteraceae is one of the largest in the number of representatives and form highly specialized group of dicotyledons. The subclass has 5 orders, 13 families, about 1400 genera and about 30 000 species. At the same time the family includes more than 90 % of the genera and species of the subclass. The annual and perennial grasses dominate; shrubs, trees, woody vines are found as a rare exception, in the tropics or subtropics. The corolla of the representatives of this class is always gamopetalous. There is often a functional and morphological differentiation of the flowers in the inflorescences, which is especially true for members of the family Asteraceae. Ovary is lower and consists of two carpels. The order Asterales. This order includes only the aster family (Asteraceae), or Compositae. One of the largest families of flowering, numbering 1250–1300 genera, 25 000 species. Species are spreaded around the world, on all continents and in all areas. Most members of the family are perennial or annual herbs, but in the tropics there are herbaceous and woody vines, stem and leaf succulents, shrubs and trees The root system is various: a rod, in some cases, a root vegetable forms. Leaves are simple, entire or lobed, alternate. The flowers are small, gathered on a broad disk, that is a shortened inflorescence’s common bed (the anthodium), often grouped into complex aggregative inflorescences — spike, racemes, panicles, cyme or capitulums. Basis anthodium is the common receptacle, which can be concave, convex or flat mesh, pitted or smooth, bare or covered with membranous scales or bristles (modified bracts) surface. On the periphery and outside the lodge is surrounded by mutated apical leaves of inflorescence — wrapper. Leaves of involucre are situated in 1, 2 or more rows, and their tops sometimes turn into spikes or hooked bristles. In the simplest case, the involucre plays the protective role for the whole inflorescence, however, it often carries additional features. The immortelles, cat’ paws, and many other genera bracts are membranous. Size of anthodiums may vary from a few millimeters to more than 10 centimeters, and the number of flowers therein ranges from 1 to 1000 or more (in cultivated sunflower forms). The anthodium is quite often a highly specialized flowers, occupying a specific position in the inflorescence. Calyx always lays in 5 tubercles, and usually few or many fuzz develops, or bristles and turns into a crest. Asteraceae have the following types of flowers: Tubular — corolla is actinomorphic — limb from 5 jags.

95 Infundibuliform — corolla is zygomorphic. False ligulate — corolla is zygomorphic. Semiflosculous — corolla is zygomorphic, limb is presented by ligule that is formed by fusion of five petals Initial actinomorphic corolla is typical for tubular flowers. In this case, the petals are fused into a tube, and their tops form a short limb of 5 jags. Corolla is zygomorphic in other types of flowers. At the edge of the funnel flowers, that are typical for, for example, cornflowers (Centaurea sp.), Zygomorphic limb forms a wide, large funnel from several unequal shares. The flowers of this type are often sterile. Corolla’s limb in bilabiate flowers is represented by two lobes that were formed by two and three tops petals. At the edge of false ligulate flowers labellum, that was formed by two bending jags, is completely reduced. In the flower corolla’s limb represented by one large petaloid ligule that is formed by the fusion of the five petals. The ligule of the corolla is almost always carries on the top 2, 3 or 5 jags (the number of lobes that was involved in its formation). The mentioned types of flowers have many transitional forms and with varying frequency occur in the family, combining in different variants. Depending on the type of flowers that are included in the inflorescence, this family subdivided into 2 subfamilies: Subfamilies Tubиliflorae (Asteroideae) — flowers are tubular, funnel- shaped, lingulated Tussilago farfara L. — the Coltsfoot Senecio platyphylloides L. Calendula officinallis L. — the pot marigold Helianthus annuus L. — the common sunflower Arnica мontana L. — the wolf’s bane Bidens tripartitа L. — the Three-lobe Beggarticks Inula helenium L. — the elfwort Matricaria chamomilla L. — the camomile Artemisia absinthium L. — the wormwood Gnaphalium uliginosum L. — the uliginose Achillea millefolium L. — the yarrow Helichrysum arenarium L. — the sandy immortelle Tanacetum vulgare L. — the tansy Subfamily — Liguliflorae (Lactucoidеae) flowers are lingular Cichorium intybus L. — the chicory Common Taraxacum officinale L. — the dandelion Raponticum carthamoides L. — the Rhaponticum Centaurea cyanus L. — the cornflower

96 LECTURE № 11 SYSTEMATIC REVIEW OF THE CLASS LILIOPSIDA, SUBCLASSES ALISMATIDAE, LILIIDAE, ARECIDAE

Class Liliopsida, or Monocotyledones. The class monocotyledons is divided into 4 subclasses and includes 37 orders, 122 families, about 3100 genera and about 63 000 species. Monocotyledonous is separated from bipartite at the dawn of the evolution of flowering plants, but the most primitive of them still retain some similarities. Among modern bipartite the greatest similarity is observed in representatives of the order Nymphaeales, however, due to the high specialization Nymphaeales can not be regarded as direct ancestors of monocotyledons. As Nymphaeales and the ancient monocotyledons appear from the more primitive moistureloving herbaceous terrestrial dicotyledonous. Subclass Alismatidae. There are 500 species and 56 genera that are grouped into 18 families belonging to 11 orders. Almost all of them are marsh, coastal, swimming, or even entirely submerged herbs. They preserve apocarpous gynoecium and extremely primitive vascular system, consisting almost entirely of tracheids. Representatives of the superclass are closest to primary monocotyledons. The order — Butomales. The order includes only 1 family Butomaceae with 1 species — Butomus umbellatus L., that can be found almost on the hole territory of Russia, except the Arctic and polar regions. This is a large perennial coastal aquatic plant forms umbellate inflorescences with beautiful light-pink flowers. Actinomorphic perianth consists of 6 leaves of similar structure. Androecium includes nine stamens and gynoecium formed only 6 carpels that are fused at the base. *Р3+3А6+3G3+3. Fruit is the polythalamous follicle. Thick starchy rhizomes of the susak are edible in baked form. The roots and the seeds of this pretty ornamental plant are used in medicine, and long narrow leaves are suitable for weaving. The order Alismatales. It includes 2 families. The family Alismataceae includes 13 genera and about 100 species. Alismataceae is a rosette perennial or annual, aquatic or terrestrial herbs with a very diverse leaves. The flowers are collected in raceme or panicle, actinomorphic, bisexual, with a double perianth. Perianth of 3 greenish sepals and 3 colored petals. There are 6 stamens, gynoecium is apocarpous, consists of numerous (rarely 6 or 3) carpels, arranged circularly or spirally, sometimes fused at the base.*Са3Со3А6 or А∞G(∞). The fruit is a poly nut, one-seeded fruits have filmy fringe, that help them to be spreaded by the water. The representatives: alisma common (Alisma plantago-aquatica L.) and Sagittaria sagittifolia L., that are growing on the banks of ponds, small bogs, and sometimes in ditches or long dry up the puddles. Rhizome is used in folk medicine as a diuretic.

97 The subclass Liliidae. This is the largest subclass of monocotyledons, including 21 order, 96 families, bringing together about 2700 genera and 56 000 species. Separate groups of the subclass reached a very high level of evolutionary development. In most cases, they form a flower with 3 sepals, 3 petals, 6 stamens and 3, in varying degrees fused, carpels. However, flower and vegetative organs of representatives of the subclass can vary within very wide limits because of the specialization. In most cases they are terrestrial plants, often passing to the ephemeral life. The order Liliales. The order brings together 9 families, the largest and most famous of which are family Melanthiaceae, Iridaceae and Liliaceae. The family Melanthiaceae includes 47 genera and about 400 species. This are perennial rhizomatous, bulbous or bulbotuberiferous herbs. Actinomorphic bisexual flowers are often collected in raceme inflorescences, rarely single. Sepals free or partially fused into a short tube. There are usually 6 stamens, and gynoecium consists of nearly 3 free carpels. Fruit is aggregate or capsule. They are distributed everywhere, many of them are cultivated as ornamental plants. The representatives: hellebore of Lobel (Veratrum Lobelianum Bernh.) and the autumn crocus (Colchicum autumnale L.). All parts of the plant are poisonous, and can cause serious poisoning of herbivores animals. Some alkaloids that can be allocated from the melanthiaceae (crocus), especially colchicine is the mitotic poisons that affect the normal chromosome segregation during cell division. The drugs from these plants are able to inhibit the growth of a tumor, but their use prevents high overall toxicity. Rhizomes of hellebore are used in veterinary medicine as an antiparasitic agent. The family Iridaceae includes 70 genera and 1500 species. It is presented by perennial herbs with creeping fleshy rhizome, bulbs or corms. The leaves are alternate, double-row, sword-shaped, sessile, linear. Flowers are bisexual, zygomorphic or actinomorphic, large, brightly colored. Perianth are double 3-membered (6 petals), the petals and sepals are often free and vary in shape, size and color, sometimes fused into a tube. There are 3 stamens, gynoecium consists of 3 carpels, inferior ovary. *Р3+3А3G(3). Fruit is polyspermous capsule, dehiscing along midrib of each carpel. The seeds have appendages, eaten by ants(they spread the seeds). Sometimes the seeds have samara and they are dispersed by wind. The representatives: yellow iris (Iris pseudаcorus L.), rhizome contain essential oils, flavonoids and etc., and are the part of the collection by the prescription M. N. Zdrenko; I. sibirica L., gladiolus imbricated (Gladiolus imbricatus L.). Currently, many ornamental varieties of hybrid iris are crocuses, or the crocuses (Crocus sp.), freesia (Freesia sp.), Gladiolus sp. The family Liliaceae contains 10 genera and about 500 species. Bulbs are typical for all representatives, these plants are often transferred to the ephemeral life. Their thicker roots are able to pull the bulb in the soil. Leaves are alternate,

98 often in a basal rosette, entire. The floral stem, often called flower arrow, leafless and leafy and carries one or more flowers in raceme inflorescence. The flowers are large, brightly colored, bisexual, actinomorphic. Perianth is 3-membered, corolliform, rarely calycine, the free or gamopetalous. There are 6 stamens (in two rounds). Ovary is superior, that is formed by 3fused carpels. *Р3+3А3+3G(3) or *Р(3+3)А3+3G(3). Flowers are pollinated by insects. Fruit is a dry capsule 3-6-herdal with numerous, usually winged seeds spreaded by the wind. The representatives are white and brindle Lilies (L. candidum L., L. tigrinum Ker-Gawl.). The order Amarillidales includes 15 families. The family Asphodelaceae has about 50 genera and nearly 1500 species. The family is represented by perennial herbs and shrubs. Rhizome often has thickened roots. Leaves often radical. Inflorescence is the alar raceme from yellow and orange tubular flowers that are located on almost leafless stalk and that have 6 perianth leaves, 6 stamens and superior ovary of 3 fused carpels. Filaments are usually densely covered with long fuzz. The fruit is almost always capsule. The most famous representer of the family is aloe vera, or century plant (Aloe arborescens Mill.), is often grown at the home. Perennial leaf succulent. Leaves are fleshy, sword-shaped. All raw materials are biogenic stimulators. Some drugs, that contain aloe juice, are widely used in medicine products. The family Hyacinthaceae includes 40 genera and more than 900 species. Bulbs and the transition to the ephemeral way of life (with a more or less prolonged period of rest) are typical for that family. Like the Liliaceae, roots of this plants can pull bulb into the soil. The flowers are collected in a receme on a leafless stalk. They are large, brightly colored, bisexual, actinomorphic, with 6 perianth leaves, 6 stamens and a superior ovary of 3 fully intergrows carpels. Fruit is dry or fleshy capsule with numerous seeds that provid with juicy appendages. Many of representatives since antiquity are favorite early-flowering plants. The representative of the family — Siberian squill (Scilla sibirica Andrews), the usual early-flowering plant deciduous forests. The family Alliaceae includes 30 genera and about 700 species. The representatives of this family are almost always have a bulb or rhizome with short socket lanceolate or linear, flat or tubular leaves. Flowers are bisexual, actinomorphic. They usually have 6 perianth segments, 6 stamens and the superior 1 or 3 locular ovary, consisting of 3 fused carpels. The flowers are collected in an umbel inflorescences at the top of leafless, rarely deciduous flower stalk, which at the beginning of the development lies in membranous spathella. The fruit is dehiscing triangular capsule with a relatively small number of large seeds. The presence of volatile sulfur oils, defining their smell is typical for this family. Oils have a strong bactericidal effect. The bulbs and

99 leaves contain a number of vitamins. All this determines the widespread use of many onions in medicine. Representatives are onion (Allium cepa L.) and garlic (A. sativum L.). The order — Asparagales. There are 8 families. Representatives are mostly rhizomatous perennial herbs. The flowers are single or in racemose inflorescences, nearly always bisexual, actinomorphic, with 6 free or fused perianth segments, with 6 (rarely with 4) stamens, and the top 3-locular ovary formed by 3 fused carpels. The fruit is almost always juicy, berry-like. The family Convallariaceae brings together 23 genera and about 230 species. Lily of the valley (Convallaria majalis L.) is the most common in the middle zone. The plant is attractive for its white fragrant flowers, the presence of glycosides, that effectively stimulate the heart. Other conventional representatives of our flora: Solomon’s seal (Polygonatum sp.), True Lover's Knot (Paris guadrifolia L.), subterranean parts contain alkaloids, all parts of the plant are poisonous, they are used in homeopathy. The family Asparagaceae includes 2 genera and about 300 species. This family is represented by lianoid, tuberforming, rhizomatous herbs whose leaves are replaced by linear or needle-like cladodes. The Representatives: Asparagus officinalis L., perennial herbaceous plant with erected, bared, branched stem. Cladodes are located on 3–6, thin, straight, thread-like, leaves are scale-like. Flowers are small, dioecious. Perianth of 6 petals, androecium contains 6 stamens, gynoecium of three carpels. The fruit is a globular berry. Aerial parts are used for diseases of the kidneys, heart, bladder. Fruits and seeds are also widely used. The plant is used as vegetable delicacy, that is very rich in lysine. The family Dracenaceae includes a number of ornamental plants covered soil. Dracaena (Dracaena sp.) are weakly branchy small trees with a rosette of large leaves at the tops of the trunks. Sansevery (Sansevieria sp.) was popularly known as ―pike light‖, because of the striated leaves. The order Cyperales. The family Сyperaceae has about 100 genera and over 4000 species. They are common almost everywhere and in the evolutionary line continue anemophilous subclass lily. They are herbaceous perennials, rarely annuals. Sedge are exclusively wind-pollinated plants. Stems, at most, are three-rowed with triangular flat gramineous-like stiff leaves. Leaves are vaginal, with narrow flat rim or cilia. The flowers are small, inconspicuous, bisexual or unisexual, clustered in an inflorescence spike, rarely in the head. Some spikes, that often bear flowers of only one gender, form a complex inflorescence. Perianth reduced to 6 or 3 scales or modified up to bristles or hairs. Three stamens have long drooping thread. Superior ovary formed by 2–3 fused carpels and contains one ovule. Female flowers are enclosed in the so-called sack, which is a bract, inosculating its edges in cone-shaped formation, the open top, (with completely reduced

100 perianth). The flowers are located by 1 in the axils black or brown nondescript axillary scales. Fruit is triangular nut enclosed in an expanding bag. Vegetative reproduction occurs at the expense of rise strongly branching rhizomes. That’s why many of them are malignant weeds. There are a lot of well-known and common herbs: reeds forest (Scirpus silvaticus L.) and S. lacustris L., various types of cotton grass (Eriophorum sp.). This moisture-loving perennial rhizomatous grasses, many of which are edificators of extremely moist cenoses. The order Poales. There is only 1 family in this order. The family Poaceae, or Gramineae occupies a special position, which is defined by the economic value and their big role in the majority of plant groups. Cereals is one of the largest families of flowering plants, numbers 650 genera and at least 10 000 species, have colonized all the habitats that are available to higher plants. Mostly they are herbs that are not capable for secondary growth. There are a lot of annuals among cereals, but by rhizomatous perennials dominate. The branching of cereals, that determines the appearance of a species, is carried near the base, in the so-called zone of tillering. The stem of almost all members of the family is a straw, segmented at the nodes and usually hollow in the internodes. The leaves are divided into sheathing leaves and linear plate with parallel venation. There is usually webbed outgrowth in its basis, that often mutates into a number of hairs. This is called the tongue or ligule. Leaf sheaths protect the base of internodes, that retain the ability to intercalated, or intercalary, growth for a long time. The flowers are bisexual, collected in a variety of complex, botryoidal inflorescences — head, raceme capitulum or compound spikes. Spikelets are the basis of all of them (characteristic elementary inflorescences of the family). Each spike may contain from 1 to 5 flowers. A typical many-flowered spike consists of a rachis, which is located near the base of the superior and lower sterile spikelet chaff, often ending with setiform arista. Flowers are placed above the chaff. The number of the flowers (1–30) is an important sign of a systematic. Each flower is located at the top of an independent two-fold axis, at its base the superior and lower floral scales are located. Chaff is modified leaves by its origin. 2 (rarely 3) small colorless scales or lodicule that are vestiges of the tepals, are usually located above paleas along the axis of the flower. Spicate, floral chaff as well as lodicules in the flower of individual species may be partially or completely reduced. In most cases, a flower has 3 free stamens. The superior unilocular ovary contains a single ovule and is formed by 3 fused carpels. The column ends with two, rarely three feathery stigmas. Flowers are pollinated by the wind. Fruit is a caryopsis, whose membranous pericarp densely adjoins to the seed, sometimes merging with the seed coat. Vegetative propagation by

101 means of rhizomes creeping, rooting shoots or bulbillate brood gemmas that are formed from modified sterile spikelets, is of great importance to the family. Representatives are the wheat (Triticum sp.), wheat starch from the grain is used in a dusting powders and ointments; рiс (Oryza sp.), corn is also a source of starch (Zea sp.), corn silk are used as cholagogic, choleretic and hemostatic remedy; barley (Hordeum sp.), caryopses are used as a source of malt, which is used for bronchitis; rye (Secale sp.), oats (Avena sp.), the decoction of the groats is used in exhaustion and as a laxative, straw is included in the tea for diabetes; the sorghum (Panicum sp.), the millet (Sorghum sp.) and others. The order Orchidales is contains one family, therefore the characteristic of the order coincides with the characteristics of the family. The family Orhidaceae is the largest family among monocots, and among the flowering plants can be compared with the Asteraceae. Orchids are represented by perennial herbaceous plants, terrestrial epiphytic forms and plants passed to the saprophytic way of feeding. Orchid leaves are simple, sometimes fleshy, round, elliptical, lorate sometimes lobed, usually green, sometimes with purple spots. The flowers are single, usually collected in inflorescence raceme with very short peduncle, transforming into the inferior ovary, there are also spike or umbellate flowers. Flowers are actinomorphic, with a double perianth consisting of 3 sepals and 3 petals, but with the zygomorphic androecium: the inner circle has 2 stamens, the outer — 1, all of them rooted to the gynoecium, created a ginostemy or column. The gynoecium of 3 fused carpels, ovary is inferior, the fruit is a capsule. Zygomorphic flowers appear later, their calyx is painted, one of the petals (labellum) is sharply distinguished from all other parts of the flower by size and color and form an embarkation platform for insects. There are 2 stamens and the third is visible in the form of staminode. All changing signs of the flowers associated with a narrow specialization to pollination. Pollen of most orchid is glued into a solid mass in the anther that is called pauline, then it released entirely and all falls on the stigma of another flower. Representatives of orchids in our flora are a lady’s-slipper orchid, one of the species of lady’s slipper. It is a large plant with wide elliptical green leaves and 1–2 large bizarre flowers. The yellow lip stands out from the truncated four of the perianth segments in the flower, one of which has a bipartite tip. Dactylorhiza speckled (Dactylorhiza maculuta (L.) Soo), green-winged orchid (Orhis morio L.), Platanthera bifolia (Platanthera bifolia (L.) Rich.), their root- tubers (salep) were used as a coating remedy. The subclass Arecidae. It includes about 4 orders, 7 families, 344 genera and about 6500 species. They are characterized by a gradual simplification of the flower, that is similar to the structure of weakly specialized representatives of the order Liliales. This process is combined with the formation of complex inflorescences, with strongly developed coating a leaf or spathe. There is

102 a transition to the secondary wood forms (palms), to the epiphytic (Araceae) and freely floating lifestyle (Lemnaceae). The order Arecales. This order includes only one family. The family Arecaceae includes about 210 genera and more than 300 species. These plants are widely distributed in a tropical and subtropical areas, forming a part of forest communities and other groups. Most palms are woody plants that do not have secondary growth and usually do not branch, rarely they are woody vines or almost stemless rosette or with greatly shortened stem plants. Leaves are petiolate, folded, pinnately or fan dissected, rarely entire, collected at the top of the trunk and with the petiole reache 10–15 m length. The flowers are small, actinomorphic or slightly zygomorphic, dioecious, collected in large, highly branched panicles. There are 6 leaves in a perianth. There are 6 stamens. The superior ovary is formed by 1–3 free or fused carpels. Stigmas are sessile or subsessile. Ovules located by 1 in each carpel, and 2 of 3 carpels often do not develop during the ripening of the fruit. *Р3+3А3,6,9,∞G(3) or 3. Fruits are monospermous and form a dry or fleshy drupe with endocarp that is surrounded by an expanded perianth at the base, rare berry- like. The fruits are used to obtain oil or can be be used as food. Representatives are date palms (Phoenix dactylifera L.) and coconut palms (Cocos nucifera L.). The order Arales. The order includes 2 families The family Araceae includes 110 genera and more than 1800 species, distributed mostly in moist tropical forests. They are epiphytes, herbaceous or lignified lianas, rhizomatous moisture-loving herbs or tuber forming ephemeroids. The most common representatives are calla (Calla palustris L.) and calamus (Acorus calamus L.) in the temperate zone — they are typical marshy or coastal-aquatic plants that are involved in the formation of quagmire. The leaves are alternate, petiolate with a leaf blade of various shapes. Flowers are actinomorphic, inconspicuous, with a simple 4–6 membered perianth or baretightly spiral gathered in a spadix, having at the base, brightly colored spathe of various shapes and structure. The spathe often forms a tube or trap for pollinating insects that are attracted by the smell or the temperature of the inflorescence. There are 4–6 stamens free or fused in sinandries. The flowers are unisexual, the male occupy the top of the spadix, and the female — lower. Superior 1–3 locular ovary is formed by 2–3 fused carpels or has 1 or more ovules in the locule. *Р0А6G(1) (the calla). *Р6А6G(3) (calamus). Fruits are monospermous or polyspermous berries that are collected in the infructescence. Medical raw material is a calamus rhizome, which is used as a decoction as an aromatic bitterness, and it is a member of various combination herbal medicinal products. A decoction of calla rhizomes is used as expectorant, antifebrile, diuretic remedy.

103 The family Lemnaceae finish hydrophilic line of this order. It includes 6 genera and 30 species, distributed everywhere. All the representatives of free- floating or partially submerged herbaceous plants, vegetative body which turned into a tiny plate of various shapes, that is called fronds. The roots are absent or poorly developed and do not reach the ground. Vegetative and generative gemmas laid at one of the poles. The inflorescence consists of 1–2 male and 1 female flowers. Perianth is absent, there is 1 stamen in the male flower, rarely there are 2 stamens. gynoecium is syncarpous with 1–4 ovules. The saccate fruits with few seeds. Representatives: species of duckweed — lesser duckweed (Lemna minor L.), duckweed ternate (L. trisulka L.), spirodela (Spirodela polyrhiza L.).

104 LECTURE № 12 BASES OF ECOLOGY AND GEOGRAPHY OF PLANTS

Ecology of plants studying relationships of plants with the environment, each other and other organisms. Habitat — a set of external factors that directly or indirectly influence on its condition, development, growth, reproduction, etc. The habitat consists of many elements of organic and inorganic nature, some of which may be necessary for the body, others are indifferent, and others — are harmful. Environmental factors are the elements of the environment that plants respond to adaptive changes. They are complex in nature. Existence conditions are the set of environmental factors, without which the organism can not exist. The classification of environmental factors: 1) abiotic: – edaphic (chemical and mechanical composition of the soil, its temperature, salinity, moisture, etc..); – climatic (temperature, light, air humidity and its movement, etc.); – orographic (relief); 2) biotic — impact of living organisms; 3) anthropogenic — human impact. The edaphic factors — complex of factors related with soil. Moisture of the soil, amount of soil water that is available to plants. The absolute water content in the soil depends on the amount and precipitation, the level of the shallow water, the soil particle sizes. Some of the water so strongly kept by the soil particles, so it is not available to plants. The absorption of water can be constrained by the low pH, high salinity or low temperature of the soil. Therefore, the absolute water content in the soil is not always characterize moisture availability of plants. The pH of the soils is from 3.0 to 9.0, and even to 10.0. Most plants prefer a neutral, a slightly acidic or slightly alkaline reaction of the soil solution (pH 6.5–7.5). Ions of hydrogen, aluminum, magnesium influence on acid soils and ions of sodium and hydroxyl — on alkalinity. The processes of mineral nutrition of plants are dependent on pH. Protoplasts of root cells of most plants are damaged At a pH of 3 or > 9. Fe, Mn, phosphate and some trace minerals bound into poorly soluble compounds in alkaline soils, therefore, plants can not obtain them in sufficient quantities. Concentration of ions Al3+ harshly increases At pH < 4–4.5, that are highly toxic to plants. Peat-bog and forest podzolized soils are very acidic (pH 3.0–3.5). Most of sod-podzolic soils are acidic (pH 4.0–5.0). The north black soils are slightly acidic. Neutral black soils and brown soils of dry steppes. Strongly calcareous semi-desert gray soils, alkaline soils and salt licks are alkaline.

105 Acidophiles are plants of acidic soils. Acidophilic plants are blueberries, lily of the valley, the vernal grasses, heather, mosses and others. Some of the cultural types of tea, flax, lupins, potato, timothy are acid resistantand. For lupine and potato pH 4.0–5.0 are considered to be optimal. The mother-and-stepmother, red clover, brome, anemone, Corydalis, thoroughwax, leek, common vetch and others belong to the plants of alkaline soils. Beets, alfalfa, clover can not tolerate acidic soils, but prefer neutral or slightly alkaline with a pH of 7.0–8.0. The specific and varietal reaction of plant clearly reveal in cultural forms. The chemical composition of the soil requires a certain adaptation of plants. For most plants readily soluble chlorides and sulfates are toxic, cause profound disturbances in the metabolism, accompanied by the accumulation of abnormal products or of incomplete transformation. Therefore, in saline soils can be found only those species that are resistant to the protoplasm of these salts — halophiles. Some plants (nettle, henbane black, dope ordinary, etc.). are nitrophilous species. Not many species can grow on soils containing heavy metal salts of copper, lead, zinc and others. Plants resist the harmful effects of salts by the accumulation of sugars, amides, organic acids. Chemical elements of the soil have a significant effect on the biosynthesis of the active ingredients in medicinal plants. Raw materials containing cardiac glycosides, can accumulate Mn and Mo; containing saponins Mo, Va, Sg, Cu, Mn. The volume of air that contained in the pores between soil particles, which increases from clay to sand, depends on the mechanical composition. The content of humus and mineral nutrients. The soluble compounds of nitrogen, phosphorus, potassium are important, i. e. the so-called elements of fertility. The richer in humus soil, the greater the proportion of all nutrients biogeocoenose contained in the soil, the active soft humus is always more favorable than nutrient-poor raw humus. Climatic factors Water. The most important environmental factor in the life of land plants because it is a better solvent, and therefore included in all cells and is necessary for the flow of physiological processes. Almost everywhere, except for the humid tropics, the plant experienced a temporary water shortage. We differentiate atmospheric drought associated with a decrease in the relative humidity of the air, and soil-dwelling, which is observed with a decrease in soil moisture that is available to plants. Rainfall is different in the different climatic zones, and the rate of evaporation also is different. According to the characterization of water supply area can be humid (moist) and arid (droughty). There are several ecological groups of plants relative to the water: 1. Gidatofity — they are submerged or floating, have no mechanical tissue and water conducting system. Pondweed, Elodea.

106 2. Hydrophytes are water-ground plants,their development begins in the water, but in adult shoots tops protrude above its surface. They live in the intertidal zone of reservoirs, can live on a wet soil away from water bodies. They have a well-developed system of intercellular spaces and pneumatic cavities (Aerenchyma). Calamus. 3. Hygrophytes grow in moist soils and with a high humidity. Leaf blades are thin, the development of hydatode to allocate water is typical. Impatiens. 4. Mesophytes — moderate humidity. Most forest and meadow plants our latitude. 5. Xerophytes are plants of dry habitats. Among them distinguish xerophytes and succulents. Xerophytes have special adaptations for obtaining and saving water: the small leaves or reduction (Haloxylon, Ephedra); development of waxy coating, powerful and nitidous cuticle; densely pubescent from colorless and hollow hairs (reflects sunlight and creates a leaf surface area, which retains moisture); depth stomata; dense, without intercellular spaces parenchyma with lots of mechanical elements that prevent fading; deep root system (at the camel thorn to 20 m), or superficial, but in large areas. Succulents are able to store water in the bodies of plants — cacti. An interesting groups are ephemera (annuals) and ephemeroids (perennial) — plants with very short vegetation period, attributable to the rainy season (most bulbous plants of desert areas) The temperature. Life of plants is possible only in a certain temperature range °. The temperature optima vary in different plants and even in a single plants throughout the growing season. Seeds of the temperate zone germinate at 8-25°, shoot growth begins at temperatures a few degrees above 0, the lower limit of root growth — 2–5°. The temperature optima of root growth, thus somewhat lower than the optima growth of stems. That is why the roots start to grow usually the first, long before bud break, and do not stop it until late autumn. Flowering is capricious process in a narrow temperature range. Some plants (one and biannual winter) only bloom if over several weeks were subjected to low temperatures (from –3 to –13°). It need a lot of heat to ripe the fruit and open the bud. Photosynthesis can take place when the lowest temperature is 5...–8°. Conifers (spruce and pine) assimilate carbon dioxide under these conditions. There are cold-loving, or cryophilic and heat-loving, or thermophilic, types. First confined to the polar and mountainous areas as well to the cold habitats of temperate latitudes. The second type is in tropical and subtropical climates, and in the temperate zones in the well-warmed plots. Adaptation to high temperatures. To reduce the heating plants increase the reflectivity of the leaves (presence of cuticle, a waxy plaque, hairs), and the rate of evaporation. The resistance of plants to high temperatures associated with the properties of its proteins. Some algae, that can be found in hot water reservoirs, can withstand temperatures up to +85 °С.

107 Adaptation to low temperatures. Perennials have creeping shoots in the tundra zone, which are covered with snow and protects them from freezing. This are so-called ―stlantsy‖. Cushion-shaped plants are formed in mountainous areas, allowing to maintain a constant temperature. Resistance to low temperatures at the cellular level is associated with dehydration of cells and weakening of breathing, increasing in the concentration of sugars, increased osmotic pressure, and the formation of ethylene glycol and glycerol compounds which lower the freezing point. Light. Light is an important environmental factor for photosynthetic plants. Most active parts of the spectrum are orange-red, blue-violet and ultraviolet. Yellow-green and infrared light are almost not absorbed. Plants are divided in relation to the light: 1. Photophilic (heliophyte) plants of an open habitats (steppe and meadow grass). They develop normally only when there is sufficient light. They have rosette of leaves, short stem. The flowers of some heliophyte can rotate the sun (sunflower, bidens). Palisade parenchyma is well-developed in the leaves of plants of illuminated areas, the cells are small, compact, without large intercellular spaces, and with numerous small stomata. There are a lot of chloroplasts in the cells, but they are small and have little chlorophyll. Shadow loving (stsiofity) are living in the permanent shadow. They develop normally only in shading. On the felling they often die. 2. Hade-tolerant (optional heliophyte) can easily tolerate some shade (most forest plants). Adapting to the conditions of lighting is based on the qualitative difference of pigments in plants in the illuminated and shaded habitats. Chlorophyll ―a‖ is contained in a large number of plants well lighted habitats. A significant part of the chlorophyll pigments represented by ―b‖ in shadow loving plants, differing increased ability to photosorption. In the shadow loving plants are almost devoid of red light and satisfied by blue rays. The length of daylight is important for plants. Reaction to day length is called photoperiodism. Long-day plants bloom when the length of daylight hours is at least 12 hours (potatoes, wheat), short-day is 10–12:00 (hemp, tobacco). Neutral — tomatoes, dandelion. Knowledge of this reaction is important in the introduction of the plants into the culture. Biotic factors. Living organisms can have a direct and immediate effect on other organisms, and indirectly alter the intensity of illumination, the microorganisms decomposing leaf litter, increase the humus content in the soil, etc. Biotic factors are divided into 3 groups: Phytogenic is the impact of plant- dwellers zoogenic is the impact of animal and human is the human impact. The effects of a chemical nature when some higher plants, that release the products of their metabolism into the environment, inhibit the growth of

108 other, is called allelopathy. Sometimes a secondary substances released in large quantities by living plants or by the decomposition of litter into the environment. Through the soil, they can affect other vascular plants, soil animals, bacteria and fungi. The accumulation of the secondary substances of the dominant plants in the soil may lead to the exclusion of other species from the community. The relationship between plants and herbivores resulted in the emergence of a wide range of plants protective devices: spines, thorns, dense pubescence. For example, mullein (Verbascum) is covered with a dense felt hairs and remains intact on the pastures. There is a chemical protection of plants: many biologically active substances (alkaloids, tannins and other phenolic compounds) were elaborated probably in response to herbivores. The ability of plants to produce toxins and accumulate them in their tissues gives them a competitive advantage.

LIFE FORMS AS A WAY OF ADAPTATION OF PLANTS TO THE ENVIRONMENT We have considered the individual environmental factors and their impact on plants. Under the influence of these factors, there are certain morphological appearance or life form. Life forms are historical adaptation of plants to aggregate many environmental factors, to the specifics of the habita. Life form may change with the change of conditions of existence. On the other hand, various floral areas on the condition similarities climate, soil, vegetation, systematically distant from each other may form a similar species, similar or convergent lifeforms. For example, the succulent ―kaktusovidnye‖ forms in Central America presents a cactus, and in Africa — plants of the family Euphorbiaceae. In the highlands of almost all regions of the world can find a variety of cushion-shaped plants. Woody plants include 3 types: the trees — plants with developed long woody stem, continuing throughout life. Shrubs have the main trunk at the beginning of life, and then he lost among equals main axes. The main axis of shrubs is available only in the early ontogeny, and then it is replaced by the lateral axes that arise from dormant buds of the parent axis (Ericaceae, Vacciniaceae). Semi woody plants — shrubs, and dwarf semishrub. Their which are retained and the basal part is lignified of the aerial shoots and gemmas of resumption located near the soil surface. The differences between the half- shrubs and woody shrubs in height of the shoots (at half-shrubs, it is 20–30 cm and semishrubs — up to 10 cm). Semi woody plants — shrubs and semishrubs, they have the basal lignified part of the aerial shoots and their buds of resumption are located near the soil surface. The differences between the half- shrubs and woody shrubs in height of the shoots (at half-shrubs, it is 20–30 cm and semishrubs — up to 10 cm). Terrestrial herbaceous plants are divided into 2 types: herbaceous polycarpic and grassy monocarpic. Each of these includes a number of groups and subgroups.

109 Water herbs are composed of 2 types: floating and submerged grass and amphibians grass. Herbaceous plants are also conveniently divided into annuals, biennials with the development in the first year rosette of leaves and perennials. Since the life forms are adapted to experience unfavorable periods of the year, so different life forms dominate in different climatic areas. The spectrum of life forms is a reflection of the climatic conditions. The most famous system of the life forms made by Raunkiaer is based on differences in position of the gemmas of resumption or shoot tips towards to the surface of the soil (or water) during unfavorable time. There are 5 types of life forms: 1. Fanerofity. The gemmas are arranged above 30 cm above the ground. 2. Hamefity. The gemmas of resumption at an altitude of 20–30 cm. 3. Hemicryptophytes. The gemmas of resumption are located directly on the surface of the soil under the litter. 4. Cryptophytes. Their aerial shoots die back completely, the gemmas of resumption stored underground at different depths. Depending on the nature of the substrate cryptophytes divided into subgroups: 1) geophytes (gemmas of resumption are stored under the ground); 2) hydrophytes (gemmas of resumption are stored under water); 3) grass helophytes (marsh plants whose aerial shoots are above the water, and gemmas of resumption are under water) 5. Terofity are annuals that experience the unfavorable period in the form of seeds. Physionomic system of life forms: trees, bushes, shrubs, cushion plants, succulents, vines, grasses, mosses and lower plants. The bases of geobotany. The vegetation cover is the entire set of plants, forming various types of vegetation communities within any portion of the earth’s surface. Individuals of species of plants are combined with each other in certain proportions forming vegetational communities that are called phytocenosis, which were formed for a long time, when the species were chosen so that they can coexist in these environments. The study of the phytocenoses is the subject of phytocenology. Regularities of distribution of vegetational communities on the surface of the globe are subject geobotany. Each area is characterized by different vegetational communities that occupy different habitats. The phytocenosis is not an apart natural phenomenon. This is the vegetational part of biogeocoenosis. But in terrestrial biogeocenoses plants occupy a predominant position and the relationship between them determine ts whole system. Each phytocenosis is characterised by the peculiar feature of the structure (species composition, life forms spectra, spatial distribution). Association is the basic unit of ecological community on which the classification of vegetation is build. Recently, the association is allocated on

110 the floristic basis. Plant association has layering the vertical dismemberment (there are no more than three in the forest zone -). The tier stands for life forms. The name of the association — from the ruling species of tiers: fir green moss, pine cranberry. The vegetation is a set of phytocenosis in a particular area. Type of vegetation is the highest rank of classification of the vegetation that combines similar plant formations that stand out on the ruling life form. Patterns of distribution primarily associated with climate zones (latitudinal zoning) and altitude zones in the mountains (vertical zonation). Ideally, the borders of climatic zones and vegetation zones on the plains should be the same, but really they are displaced due to the varying distance from the ocean (especially continental). The vegetation of freshwater bodies, saline areas, wetlands is intrazonal. There are the following areas of vegetation: Tundras. The climate is harsh. The annual average temperature is below 0, the vegetation period is 2 months. The permafrost is almost everywhere. The vegetation consists of mosses, lichens, grasses, creeping dwarf trees and shrubs. Coniferous forests. Located in the temperate climate zone. They are evergreen, except larch, mostly homogeneous: in the northern parts of Europe — they are spruce, in the middle lane — piny, Siberian — larch forests. They are characterized by continuous moss cover. There are many pileate mycorrhizal fungi. They are often waterlogged. Deciduous broad-leaved and small-leaved forests. The growing season is 4–6 months with ample rainfall. There are mixed forests on the border with piny forests -. The predominant species are the beech, hornbeam, oak, birch, aspen, alder. Ephemeroids vegetate before leaf flushing. There are a large number of herbaceous species. The sclerophyllous forest. West Caucasus, the southern coast of Crimea, Mediterranean. Summers are hot and dry, winter is warm and humid. They are evergreen, flowering from december to april. Scrubs (maquis) are formed by dwarf oak, laurel, myrtle, oleander. Steppes. The climate is continental arid. The herbal plant communities. Cereals dominate — feather grass, fescue, crested, and others. The soils are black earth, brown, very pinguid. Deserts and semi-deserts. The climate is continental, arid. Poor vegetation do not form a canopy cover. There are a large number of ephemera and ephemeroids expressed xerophytes: wormwood, grasses, Haloxylon. The salt desert there are salicornia, halophytes. Intrazonal vegetation. High moor bogs — sphagnum bogs (moisture precipitation accumulates without runoff). Sphagnum, cranberry, sedge, sundew. Lowland — swamps with high standing of the groundwater. Hygrophilous grasses, sedges, cotton grass.

111 Modern Earth’s vegetation is composed not only of natural but artificial phytocenoses. The natural vegetation covers 65.9 % of the territory in Belarus. The predominant type of vegetation is forest (35.5 %). In the middle of the XVIII century. The forest area was more than 74 %. The formations of spruce, oak, ash forests are zonal in Belarus. Grassland ecosystems occupy 15.8 %, 11.5 % marsh and shrub 3.1 %. Geobotanical zoning is the division of the territory into regions that are homogeneous in composition and structure of vegetation, soil and orographic conditions. In Belarus there are three subzones (from north to south): 1 oak- conifer, 2-hornbeam-oak-dark coniferous, deciduous 3-pine forests.

THE GEOGRAPHY OF PLANTS (THE FLORAL GEOGRAPHY) The concept of flora. Flora is the set of species of plants that can be found in the region making up all its characteristic plant communities that plant all types of habitats. The idea of the flora does not apply generally to the species that are found in the area just as plants, bred by man. Plants, deliberately or accidentally listed by person, but wild and supporting their existance independently of human activity is the part of the flora. The most important feature of each flora is the species composition. The natural flora of Belarus totals 1737 species, and 646 of them are anthropophytes. Flora are distinguished by the wealth (the total number of species), the complexity of the composition (distribution of species by the genera and families), the peculiarities of the taxonomic composition (the presence of specific groups); by connection floras of the specific shape and composition with the complex of environmental conditions (flora of the cold and hot, arid and humid areas, and so on.); by their connection with the dominant types of vegetation (forest, steppe, desert, tundra flora and so on.); by the connection with certain elements of the relief (for example, the flora of the plains, mountains, high mountains); geographical and genetic elements; on the ―age‖ of flora, and so on. Different floras are characterized by different relations between woody and herbaceous plants. The sharp prevalence of the latter is characteristic of the Northern and Central Europe, Siberia. Among the herbaceous plants of these spaces, in turn, perennial forms are sharply dominated. The Mediterranean region is characterized by a high number of annuals — they are: ephemera (30–35 %) of the species composition. For tropical regions — trees, vines; among herbaceous plants — a lot of epiphytes. Classification of the floras reveals patterns of spatial distribution of floristic complexes of different shape, different taxonomic composition and origin.

112 Учебное издание

Гурина Наталия Сергеевна Кузнецова Ольга Анатольевна Мушкина Ольга Владимировна Волочник Мария Валерьевна

ФАРМАЦЕВТИЧЕСКАЯ БОТАНИКА ДЛЯ ИНОСТРАННЫХ СТУДЕНТОВ

PHARMACEUTICAL BOTANY FOR INTERNATIONAL STUDENTS

Учебно-методическое пособие

На английском языке

Ответственная за выпуск О. В. Мушкина Переводчики: Н. С. Гурина, О. А. Кузнецова, О. В. Мушкина, М. В. Волочник Компьютерная верстка Н. М. Федорцовой

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