CS 153: BOTANY OF CROP PLANTS Course Outline Introduction Branches of Botany Plant Classification and Nomenclature Plant Structure Seed Morphology Root: Functions, Internal Structure, Types, Modifications Stem: Functions, Types, Internal Structure, Modifications Leaf: Types, Functions, Internal Structure, Leaf Venation, Phyllotaxy Reproduction in Plants Flower: Types and structure, Pollination and Fertilization Fruit Development and Types Dispersal of seeds and Fruits References 1. Botany (An Introduction to Plant Biology) 5Ed, James D, Mauseth 2. Botany for Degree Students, A. C. Dutta (Revised Edition) 3. Principles of Botany. Gordon Uno, Richard Storey, Randy Moore 4. others AA AMOAH 1 CS 153: BOTANY OF CROP PLANTS Introduction The science that deals with living objects goes by the general name of biology (bios, life; logos, discourse or science). Since both plants and animals are living, biology includes a study of both. Biology is, therefore, divided into two branches: botany (botane, herb) which deals with plants and zoology (zoon, animal) which deals with animals. Scope of botany Botany deals with the study of plants from many points of view. This science investigates the internal and external structures of plants, their functions in regard to nutrition, growth, movement and reproduction, their adaptations to varying conditions of the environment, their distribution in space and time, their life history, relationships and classification, the laws of heredity, the uses that plants may be put to and, lastly, the different methods that can be adopted to improve plants for the use of mankind. What is Botany? Provide the definition as taught in class. Branches of Botany 1. Morphology (morphe, form; logos, study): The study of the external form of the plant and how it attains it. 2. Anatomy: The study of the gross internal structure of a plant organ and how it comes about. 3. Histology (histos, tissue): The study of the detailed structure of tissues making up a particular organ of the plant. 4. Cytology (kytos, cell): The study of individual cells and their contents 5. Genetics: The study of heredity which is transmission of the characteristics of individuals from one generation to another. 6. Plant breeding: The use of genetic principles for the improvement of plants. 7. Physiology (physis, nature of life): This is the study of the various functions that plants perform. This includes all the living processes that go on in the leaf, stem, roots and other organs e.g. Food manufacturing, transport and storage, absorption of water and nutrients, transpiration. 8. Ecology (oikos, home): The study of the inter-relationships between plants and their environment. 9. Plant Geography: This deals with the distribution of plants over the surface of the earth and the factors responsible for this distribution. AA AMOAH 2 10. Taxonomy or Systematic Botany: This deals with the description and identification of plants, and their classification into various natural groups according to the similarities and differences between their morphological characteristics. 11. Agronomy: Is the branch of botany which emphasizes the application of botany into farm practices. It is concern with the culture, harvest and improvement of field crop. It includes plant breeding, nutrition, the maintenance of soil fertility and crop protection. AA AMOAH 3 PLANT CLASSIFICATION The Binomial System of Nomenclature The scientific name of every plant consist of two parts the first part designate genus and second species. This is what is called the binomial system. It was first proposed by the Swedish nationalist Carl Linnaeus (1707-1778). The specie is regarded as a group of closely related individuals capable of breeding successfully with each other but not with other different groups. Species that are different but related by descent are group together in the genus. Taxon order from high to the lowest (downwards) Kingdom Sub-kingdom Division / Phyla Sub-division Class Sub-class Order Family Genus Specie The plant kingdom consists of over 300,000 different kinds of plants and there are various schemes of classifications. A tentative scheme based on modern natural classifications, satisfying our needs, is as follows: Kingdom Plantae Sub-kingdom A. Thallophyta (plants not forming embryos) Division I. Phycophyta or Algae Sub-division: Cyanophyta or blue-green algae Euglenophyta or euglenoids Bacillariophyta or diatoms Chlorophyta or green algae Phaeophyta or brown algae Rhodophyta or red algae Division II. Fungi or Mycophyta Sub-division: Schizomycophyta or bacteria Myxomycophyta or slime fungi Eumycophyta or true fungi; Sub-division: Eumycophyta Class: Phycomycetes or alga-like fungi, Ascomycetes or sac fungi Basidiomycetes or club fungi AA AMOAH 4 Sub-kingdom B. Embryophyta (plants forming embryos) Division I. Bryophyta (plants without vascular tissues, they are valuable in soil formation) Sub-division: Hepaticae or liverworts Anthocerotae or horned liverworts Musci or mosses Division II. Tracheophyta (includes a large number of vascular plants and of economic importance) Sub-division: Psilotopsida (extinct) Lycopsida (club mosses) Sphenopsida (horsetails) Pteropsida (ferns and seed bearing plants) Sub-division: Pteropsida Class: Filicinae (Ferns) Gymnospermae (coniferous trees) Angiospermae (flowering plants) Class: Angiospermae Sub-class: (i) Monocotyledonae (monocots/ grasses) (ii) Dicotyledonae (dicots/ broad leaves) E.g. Classification of a monot and dicot Taxon Maize Groundnut Kingdom Plantae Plantae Division Tracheophyta Tracheophyta Sub-division Pteropsida Pteropsida Class Angiospermae Angiospermae Sub-class Monocotyledonae Dicotyledonae Order Graminales Leguminosae Family Gramineae Papilionaceae Genus Zea Arachis Specie mays hypogaea Note: underline the scientific names when writing or italize when typing Varieties (Cultivars) A variety is a group of rather closely related plant lower than a species. They are cultivated and recognized agriculturally as a unit. The quality of the products produced by such a group of plants e.g. nutritional difference and their yields are important characteristics of a variety that go beyond the botanical standards that are used to determine general species etc. AA AMOAH 5 THE PLANT STRUCTURE The plant kingdom is characterized by great diversity and form. Bacteria are microscopic lower forms of plant life, may be unicellular bodies or cells may be joined to take the form of a slender thread, or may be grouped into a colony or organized into a primitive structure called thallus. The higher forms are elaborate and complex. They have roots, stems and leaves. They may be tall and slender or short and bushy or trailing in growth with stems spreading over the soil surface or twining about erect plants. There are succulent or the herbaceous types whose stems die-back each season and woody types such as crops which continue to grow from year to year. Inspite of this diversity in the higher plant, a simple pattern of development occurs Fig. 1 The plant body consists fundamentally of axis which at one end becomes the root and at the other the shoot. The shoots intend is made up of leaves. The root serves to anchor the plant and to absorb water and minerals from the soil. A young stem (twig) is marked by the presence of nodes which are the point on the stem where a leaf or leaves are attached. The intervals between the nodes are called internodes. Axillary buds are usually found in the angle between the leaf and the stem. The buds may grow into branches that duplicate the structure of the shoot. The leaves are arranged on the stem in a manner that is constant for any given specie. The root bears no appendages comparable to the leaves and therefore has no nodes or internodes. The lateral roots may be regarded as branches of the root. But unlike the shoot the branches arise within the tissues of the parent roots rather than externally visible buds. AA AMOAH 6 CLASSIFICATION OF PLANTS BASED ON LONGEVITY OF AERIAL PARTS They may be classified according to length of life of the entire plant or sometimes only of its aerial portion. 1. Trees are large plants with a single stout trunk and hard, and woody branches profusely formed (except most palms). 2. Shrubs are medium-sized plants with hard and woody stems which branch profusely from near the ground so that the plants often become bushy in habit without having a clear trunk. Trees and shrubs have shoots that live for a number of years. 3. Plants in which the aerial portion is relatively short lived are called herbs or herbaceous plants. Based on the duration of their life they may be classified as a. Annuals: plants that attain their full growth in one season, living for a few months or at most for one year producing flowers, fruits and seeds within this period, e.g. rice, pea b. Biennials: plants that live for two years. They attain their full vegetative growth in the first year and produce flowers and seeds in the second year, after which they die off e.g. radish, carrot etc. (In tropical climates they behave like annuals.) c. Perennials are those plants that persist for a number of years. The aerial parts of such plants may die down every year at the end of the flowering season, but next year new shoots develop again from the underground stem after a few showers, e.g. ginger. AA AMOAH 7 SEED MORPHOLOGY Seed The seed is a complex structure composed of an embryo, seed coat and a supply of stored food. The most important of them is the embryo. Embryo is a very young plant and consist of a primary axis with the following component parts root, hypocotyls and stem, at this stage it is difficult to differentiate where the root joins the hypocotyls, but above the hypocotyls is the node of the first two leaves i.e. cotyledons referred to as seed leaves and it marks the beginning of the stem. The components remain quiescent until the seed germinates. Types of Seeds a. Endospermous Seed: The endosperm is the nutritive tissue on which the developing embryo feeds. In some seeds including almost all monocots and many dicots, the endosperm persist and serves as food for the embryo after germination, these are endospermous seed. Examples are castor bean, rice etc. b. Non-endospermous seeds are those in which the nutritive tissue is completely absorb before the seed is ripe. Examples are bean, cucurbita etc. Fig. 2 AA AMOAH 8 Monocotyledons and Dicotyledons Seeds Monocot produce seeds each with only one cotyledon, or seed leaf. The cotyledon is a modified leaf involved in storing or supplying nutrients and energy for the embryo in the seed. Dicots produce seeds with two cotyledons. When the development of seed is complete it usually dries out and enters into a period of dormancy. GERMINATION: Is the process by which the dormant embryo wakes up, grows out of the seedcoat and establishes itself as a seedling. The embryo grows by absorbing food material stored up in the cotyledons, or in the endosperm when it is present. There are two kinds of germination, epigeal and hypogeal. GERMINATION OF THE BEAN SEED When the seed of bean is placed under favourable condition of moisture, temperature, and air, various parts of the embryo begin to grow. First, the root appears and grows downward into the substrate or soil. It is described as been positively geotrophic. Root hairs which are very small and delicate extensions of the epidermal cells from the back of the root cap absorb water and nutrients and give the plant its first anchorage. Role of hypocotyl in germination As the radicle grows downwards the hypocotyl begins to grow upwards. Since the two large cotyledons and the stems are above the hypocotyl, they are pulled upwards by the curved hypocotyl as it elongates. The curved or arched growth of the hypocotyl prevents the breaking of the cotyledon as it moves up through the soil and becomes exposed to light, further elongation diminishes rapidly except in the arch itself. The underside with much less exposure to light continues to grow while the growth of the upper and exposed side is retarded with much differential growth. The hypocotyl straightens and simultaneously pulls the cotyledon out of the soil. The cotyledon spreads apart above the soil as two fleshy leaves. They acquire green chlorophyll rapidly and manufacture food until several other leaves above them develop then the cotyledon shrivel and drop off the stem this is referred to as epigeal. AA AMOAH 9 Fig. 3 Hypogeal Germination In most monocots e.g. maize and some dicots, the hypocotyl does not elongate and the cotyledons remain below the surface of the soil. The plumule produces a stem that arches until it reaches the soil surface then it straightens and grows upwards. The cotyledons in the soil finally disintegrate after the redrawal of food by the seedling e.g. garden pea (Pisum sativum) AA AMOAH 10 SEEDING DEVELOPMENT OF MAIZE (MAIZE CARYOSIS) Fig. 4 The cotyledon in maize has undergone extreme evolutionary modification. It is composed of two main parts the scutellum which is food absorbing organ and the coleoptiles which is a protective cap over the plumule. During germination the first organ to emerge is the radicle which breaks through the coleorhiza and then through the tip of the seed. Very soon seminal roots appear at the scutellar node i.e. junction in the embryo between the radicle and the plumule. The primary root system formed from the radicle never becomes large and may be temporally. These primary roots are supplemented by a stronger secondary root system adventitious in origin which develops from the lower nodes of the stem. Emergence of the corn seedling is brought about by elongation of the mesocotyl. The mesocotyl is regarded as a combination of the hypocotyls and of the cotyledonary tissues that originally connected to the scutellum and the coleoptiles. The elongation of the mesocotyl depends on a supply of hormones. This hormone supply is adequate to stimulate the growth of the plumule and the secondary roots. When the tip of the coleoptiles breaks through the soil surface, rate of auxin production is sharply reduced by absorption of light. The growth processes are then reversed. Elongation of the mesocotyl ceases. The plumule emerges from the coleoptiles and roots develop from the first node. AA AMOAH 11 THE ROOT The root is the descending portion of the plant and is not normally green in colour, ends in a protective cap known as root cap, bears unicellular hairs and has no nodes or internodes. The root performs four key functions for the plant. A) It is the site for absorption of water and nutrients. B) The root anchors the plants and gives it stability C) The root serves as a translocation system for absorbed water and nutrients D) In some plants the roots serve as a site of storage for accumulated carbohydrate. ROOT STRUCTURE The root tip which is one of the plants epical meristem is covered and protected by a thimbleshaped structure called Root cap. New root cap cells are formed constantly in the inner of the cap. Cells of the root cap can secrete large amounts of mucigel, a slimy substance containing sugars, enzymes and amino acids that lubricates roots as they push between soil particles. Mucigel also protects roots from drying out, makes it possible for minerals, along with water in the soil, to be better absorbed by roots. Due to the impact of the hard soil particles, the outer part of the root cap wears away and newer cells formed by the underlying growing tissue are added to it. Zone of Cell Division: This is the growing apex of the root lying within and a little beyond the root-cap and extends to a length of one to a few millimeters. It consists mainly of the apical meristem of the root, where new root cells are produced. These cells divide every 12 to 36 hours; in some plants the meristem produces almost 20,000 new cells each day. Some of the newly formed cells contribute to the formation of the root-cap and others to the next upper region. Zone of Cell Elongation: This lies above the meristematic region and extends to a length of a few millimeters (1 to 5mm, or a little more). The cells of this region undergo rapid elongation (primarily by filling their vacuoles with water) and enlargement, and are responsible for the growth in the length of the root by shoving the root-cap and apical meristem through the soil. Zone of Maturation: This region lies above the region of elongation and extends upwards. Externally, often extending to a length of a few millimeters and sometimes a few centimeters, this region produces a cluster of very fine and delicate thread-like structures known as root-hairs. Root hairs provide a very efficient contact with the soil for absorption of water and nutrients. Internally, the cells of this zone are seen to undergo maturation and differentiation into various kinds of primary tissues. Fig. 5 Root Structure AA AMOAH 12 Primary tissues: The root apex (apical meristem) is a region of tissues initiation, it gives rise to the primary meristem i.e. the protoderm, ground meristem and procambium. Fig. 6 Internal Structure of the Root A cross section through a mature dicot root gives us a look at its primary structure. A typical dicot root is surrounded by a layer of epidermis cells. The root epidermis usually either lacks a cuticle (a thin coating of wax) or has a thin cuticle that does not significantly affect water absorption. Xylem and phloem cells are packed together in the center of the root. Cortex cells constitute the majority of cells in the root and often store most of the starch. In many plants, the outmost layer(s) of the cortex is a protective layer called the hypodermis, with cells lined with suberin, a waxy substance that is impervious to water. Fig. 7 Most of the cortex consists of thin-walled, starch-storing parenchyma cells separated by large intercellular spaces that can occupy as much as 30% of the root’s volume. The innermost layer of the cortex is the endodermis. Cells of the endodermis are packed tightly together and lack intercellular spaces. Furthermore, four of the six sides of each endodermis cells are impregnated AA AMOAH 13 with a Casparian strip made of lignin and suberin and arranged similarly to a rubber band around a rectangular box. If endodermis cells are likened to bricks in a brick wall, then the Casparian strip is analogous to the mortar surrounding each brick. The Casparian strip redirects the inward movement of water and nutrients as they flow through the endodermis; water and dissolved minerals must pass through the cell membranes of endodermis cells to reach the vascular tissues in the center of the root, the stele. Fig. 8 The stele of a root includes all of the tissues in the middle of the root and consists of the pericycle, vascular tissues (xylem & phloem), and sometimes a pith. The outermost layer of the stele is the pericycle, a layer of thin-walled meristem cells one to several cells thick. It’s important because it produces branch roots. The earliest sign of branch-root formation is cell divisions in the pericycle. Soon thereafter, a root cap and primary tissues form. The branch root then forces its way through the soil. The vascular tissues of branch roots link with those of the parent root. Fig. 9 Inside the pericycle is the root’s vascular tissue. Roots of most dicots and some monocots (e.g. wheat and barley) have a lobed, solid core of xylem in the center. Phloem is found between the lobes of xylem. The roots of many monocots and few dicots have a ring of vascular tissue surrounding pith made of parenchyma cells. AA AMOAH 14 Fig. 10 Note arrangement of vascular bundles in root as compared to the stem. Note the arrangement of xylem to the phloem (i.e. alternate arrangement instead of all being on the same radius) TYPES OF ROOTS The primary root and its branches form the tap root system of the plant. The primary or tap root normally grows vertically downwards to a shorter or longer depth, while the branched roots (secondary, tertiary, etc.) grow obliquely downwards, or in many cases spread horizontally outwards. This is common in cone-bearing trees and dicots. When a primary root is not dominant in its growth and it is superseded by the growth of its branches or fine roots arising out of the lower nodes of its stem, the plant has a fibrous root system as occurs in most monocots (including grasses). Fig. 11 ADVENTITIOUS ROOTS Roots that grow from any part of the plant body other than the radicle are called adventitious roots. They may develop from the base of the stem, replacing the primary root or in addition to it, or from any node or internode of the stem or the branch, or even from the leaf, under special circumstances. Fibrous and Foliar Roots are forms of adventitious roots. AA AMOAH 15 MODIFIED ROOTS Aerial Roots: these are adventitious roots because they grow from the stem. They serve several functions including the following, some are photosynthetic, some are breathing roots (pneumatophores) these roots have lenticels in the outer tissues for gaseous exchange, aerial roots for additional support e.g. prop roots in maize, climbing roots in betel (Piper betle), buttress roots in kapok (Ceiba). Storage roots: roots of most plants act as storage organs without being modified in any way. In carrot (Darcus carota) and beet root (Beta vulgaris) the tap root is modified by being greatly swollen. In both plants the storage organ starts to form at a very early stage and the top part is the swollen hypocotyl. Root tubers: it’s a swollen lateral or adventitious root. The edible tuber of cassava (Manihot esculenta) and sweet potatoes (Ipomoea batatas) are both swollen adventitious roots. AA AMOAH 16 THE STEM Characteristics of the Stem The stem is the ascending portion of the axis of the plant, developing directly from the plumule, and bears leaves, branches and flowers. When young, it is normally green in colour. The growing apex is covered over and protected by a number of tiny leaves which arch over it. The stem often bears multicellular hairs of different kinds. It branches exogenously and is provided with nodes and internodes which may not be distinct in all cases. Leaves and branches normally develop from the nodes and buds. When the stem or the branch ends in a vegetative bud it continues to grow upwards or sideways. If, however, it ends in a floral bud, the growth ceases. TYPES OF STEMS There are varieties of stem structures adapted to perform diverse functions. They may be aerial or underground. Aerial stems may be erect, rigid and strong, holding themselves in an upright position, while there are some too weak to support themselves in such a position. They either trail on the ground or climb neighbouring plants and other objects. 1. Erect or Strong stems e.g. i. Caudex is an unbranched, erect, cylindrical and stout stem, marked with scars of fallen leaves as found in palms. ii. Culm is a jointed stem with solid nodes and internodes as found in bamboo. iii. Some herbaceous plants, particularly monocotyledons, have no aerial stem. The underground stem in them produces an erect unbranched aerial shoot bearing either a single flower or a cluster of flowers; such a flowering shoot is called scape, as in onions. 2. Weak Stems: these are commonly of three kinds A. Trailers: trailers are those plants whose thin and long or short branches trail on the ground, with or without rooting at the nodes. When such plants lie prostrate on the ground they are said to be i. Prostrate or procumbent, e.g. Oxalis. ii. When the branches of such plants after trailing for some distance tend to rise at their apex they are said to be decumbent e.g. Tridax. iii. When the plants are much branched and the branches spread out on the ground in all directions, they are said to be diffuse e.g. boerhavia. B. Creepers: Are weak-stemmed plants with long or short branches creeping along the ground and rooting at the nodes. A creeping stem may be a runner, stolon or sucker according to its varied nature. C. Climbers: climbers are those plants that attach themselves to any neighbouring object, often AA AMOAH 17 by means of some special devices e.g. pea, yam etc. Nodes and Internodes: The place on the stem or branch where one or more leaves arise is known as the node, and spaces between two successive nodes is called the internode. Bud: A bud is a young undeveloped shoot consisting of a short stem and a number of tender leaves arching over a growing apex. In the bud, the internodes have not yet developed and thus, the leaves remain crowded together, forming a compact structure. The lower leaves are older and larger than those higher. The normal position of a bud is at the apex of the stem or branch and in the axil of a leaf. The bud in the former case is known as the terminal or apical bud, and in the latter as axillary bud. Sometimes some extra buds develop by the side of the axillary bud. These are known as accessory buds. Fig 12 Functions of the Stem: Main purpose is to bear leaves and flowers and spread them out on all sides for proper functioning. Support of the branches which push forward the leaves and flowers Conduction of water, mineral salts Storage of water and food AA AMOAH 18 INTERNAL STRUCTURE OF THE STEM Fig 13 MONOCOT STEMS The structure of the monocot stem differs from that of dicot and conifers in two main ways. 1. Vascular tissues are usually organized into separated bundles and these are scattered throughout the stem instead of being in the cylindrical arrangement, as a result, no distinction can be drawn between the pith and cortex in some monocots such as bamboo, wheat and other grasses. The internodal regions are hollow, but even in these plants the vascular bundles are irregularly arranged. 2. All the cells of the procambial strands mature into xylem and phloem. The cambium is therefore absent since they lack a lateral meristem. The tissues of a monocot stem are primary in origin. Growth in Vascular Plants Two kinds of growth occur in vascular plants. These are primary and secondary growth. Primary growth is initiated in the apical meristem in the shoot and the root. There are three sections of development Section A: includes the shoot apex where cell division is dominant but some cell enlargement and differentiation are also dominant. Section B: cell division continues but cell enlargement and tissue differentiation are more conspicuous. Section C: cell enlargement continues but, more tissues at this point have become differentiated and tissues such as epidermis, pith and cortex are approaching physiological stages. This primary growth is completed when derivative of the apical meristem has differentiated into the mature tissue which constitute the primary body. AA AMOAH 19 Tissues of the Primary Body These are the primary phloem and xylem, the cortex, pith and epidermis. These primary tissues are derived from three tissue systems which can be found below the apical meristem. 1. The dermal tissue is called protoderm which forms the epidermis 2. The meristematic vascular tissue is the procambium which gives rise to the primary vascular tissues. 3. The ground meristem produces the ground tissues which are the pith and cortex. VASCULAR TISSUE The procambium is composed of vertically elongated cells arranged in dicot and coniferous stem in a cylindrical pattern. The outer cells of the procambium strand differentiate into primary phloem and the inner into primary xylem. Between the two there are remains of a single layer of cells called the vascular cambium which does not become differentiated but remains meristematic. These cells retain indefinitely their capacity to divide to form new cells. Thus causes increase in diameter of stem. PRIMARY XYLEM TISSUES Fig. 14 The first form of primary cells is termed annular elements. The first formed or inner most of the primary xylem cells are termed annular elements because the secondary wall is laid down as rings or annular thickens. Others which differentiate later are spiral, scalariform, reticulate and pitted. All the above types of primary xylem cells don’t necessary occur in a particular plant or plant organ. PRIMARY PHLOEM TISSUES The primary phloem often includes a considerable amount of thick walled supporting elements called sclerenchyma usually in the form of elongated fibres. Food may be stored in the cortex of herbaceous stem and collenchyma cells may also occur in the cortex which is able to manufacture food because they contain chlorophyll. Collenchyma is also an important primary tissue which supplements the mechanical support function of the vascular tissue. AA AMOAH 20 SECONDARY GROWTH Secondary growth results from the activities of a lateral meristem mainly the vascular cambium. It is characterized by thickening of stem and roots and the tissues formed during such growth are called secondary tissues. The cambial cells divide and produce secondary xylem (wood) on the inner side and secondary phloem (bark) on the outside. By the end of the growing season the first growth or annular ring has been formed. Fig. 15 Secondary growth of dicotyledonous stem COMPARISM BETWEEN PRIMARY AND SECONDARY GROWTH Primary growth is responsible for increase in height and secondary growth for increase in diameter. Primary tissues make up a relatively small amount of the volume of woody stem and they play an insignificant role in the economy of the plant. In many herbaceous plants however, primary tissues constitute the bulk of the plant body. In monocot a cambium is usually absent and the plant tissues are thus primary in origin. Secondary xylem tissues include vessels, tracheids, xylem parenchyma cells and xylem fibres. The proportion of the different cell types and their arrangement vary from species to species. Hard wood contains many fibres while soft wood contains a great deal of xylem parenchyma and few fibres. *Reading Assignments (heartwood, sapwood, growth rings, sieve tubes, phloem fibres, cork (periderm), lenticels) HEARTWOOD AND SAPWOOD A large tree may have a large volume of secondary wood, but only the youngest outermost part of the secondary wood called the sapwood is alive and functioning. In the sap wood the xylem and ray parenchyma cells are living and contain stored starch and the vessels are transporting water from the roots to the leaves. The older central part of the wood is dead. This is the heart wood and it makes the most valuable timber. AA AMOAH 21 Fig. 16 GROWTH RINGS Growth rings appear as concentric circles in cross section and are usually 1 to 10 millimeters wide. Each growth ring increases the diameter of the trunk; once formed, each ring remains unchanged in size or position during the life of the tree. Because climate (especially the availability of water) strongly influences the formation of growth rings, a cross section of wood is a diary of the climatic history of a particular region. (The science of interpreting history by studying growth rings is called dendrochronology). Cambial action in tropical tree is less related to the seasons and more affected by variation in rainfall throughout the year. There are several kinds of phloem cells and this make up the inner portion of the bark just outside the cambium. Secondary phloem is composed of sieved tubes, companion cells, phloem parenchyma, and phloem fibre. SIEVE TUBES In flowering plants these are the most important cells for it is through them that vertical conduction of food occurs. They are composed of a number of sieve tube elements placed end to end. The primary walls are relatively thick and the secondary wall is usually absent. The most conspicuous feature of these cells is the presence of sieve areas which are sunken wall areas with clusters of pores. It is through pores sieve plates that food move from one sieve element to another. The functional life of the sieve tube elements is short, commonly only a single growing season. The contents become disorganized and in many plants, the sieve tubes eventually collapse as a result of pressure of surrounding cells, but in long lived monocots with no cambium they depend upon the original primary sieve tubes in the older portion of the stem. Associated with each sieve tube may be seen a smaller cell. This is the companion cell. The rest of the phloem is packed with small-celled parenchyma known as the phloem parenchyma. All the phloem elements are living, and contain various kinds of food material. PHLOEM FIBRES In herbaceous stems, these are usually restricted to the outer part of the phloem. These fibres from stem of herbaceous dicotyledons are important raw materials from which cord and textiles are manufactured. AA AMOAH 22 CORK (PERIDERM) Most woody and some herbaceous plants form a layer of cork on the outside of the stem. The cork is formed by cork cambium called phelogen which arises in the epidermis or more commonly in the outermost layers of the cortex just beneath the epidermis. The cork cambium divides repeatedly and produces a layer of cork cells to the outside and a few parenchyma cells to the inside. The cork separates the epidermis from its supply of food and water and so the epidermis dies and gradually sieves away. Later the cork cells die and become filled with waxy materials. The cork retards the loss of water from the stem tissues. It is also formed during the healing of wounds thus preventing the drying out of exposed tissues and the entrance of decaying fungi. The cork may serve as insulator and may prevent the destruction of the cambium by ground fire. LENTICEL These are structures in the bark that serve in aeration. They are composed of masses of loosely arranged cells with numerous intercellular spaces through which oxygen may penetrate to the living tissues within. It is the product of the cork cambium which locally gives rise to the cells of the lenticels instead of cork. STEM MODIFICATION The main functions that modified stems perform are: 1. Perenation i.e. surviving from year to year through bad season by certain underground stems. E.g. Subterranean stems: In many plants, the aerial parts or branches arise from underground stems; such stems usually serve as food storage sites and are important sources of food for men and animals. a. Rhizome: They are perennial stems horizontal in position. It is the main axis of the shoot with relatively short and thickened internodes, rich in accumulated food. The tip of each rhizome ultimately grows up and forms a leafy branch while lateral branches continue to grow underground e.g. ginger (Zingiber officinale). b. Stem tubers: The iris potato (Solanum tuberosum) is a good example of stem tubers. Axillary shoots or stolons are formed underground and grow out horizontally or downwards. The tip of each shoot swells with stored food, forming a tuber (tuber means a swelling). The surface of the tuber is marked at one end by a scar which was the point of attachment of the stolon and by the ‘eyes’ scattered over the surface. A single eye consists of a ridge bearing a minute scale-like leaves in the axil of which are several usually minute buds. The scale leaf and its axillary buds correspond to the leaf of a woody twig with the bud in the axil of the leaf. The portion of the tuber between consecutive eyes represents the internode. AA AMOAH 23 Fig. 17 c. Corms: Is an underground stem that is short, greatly swollen, fleshy and grows in the vertical direction e.g. Colocasia sp and cocoyam (Xanthosoma). A corm has several nodes each with a scale leaf. There is an apical bud at the end and this grows out at the start of the growing season to form a leafy plant, buds in the axils or scale leaves then form new corms which will be ready to produce new leafy shoots at the beginning of the next growing season. Fig. 18 d. Bulb: This is an underground modified shoot (rather a single often large, terminal bud) consisting of a very short convex or slightly conical stem rapped in thickened fleshy scales or leaf bases which are the site of food accumulation. New bulbs are formed at the base of a shoot and from axillary buds e.g. Allium cepa (onion) Fig. 19 Vertical cut section of an onion AA AMOAH 24 e. Suckers: These are formed from axillary buds which are formed from the under surface of the soil and grow obliquely upwards and directly gives rise to leafy shoot or a new plant. Example musa spp. (banana and plantain) which forms suckers which grow up around the parent plant and takes its place when it has fruited and die. 2. Vegetative propagation by certain horizontal sub-aerial branches spreading out in different directions. E.g. Prostrate branches: Many species that have upright shoots produce slender horizontal branches from basal node of the stem. These structures are called runners or stolons. They grow either over or under the soil surface. The term runner is usually restricted to adventitious branches and horizontal branches in the soil. In both runners and stolons nodes and internodes are present and adventitious roots and sometimes buds are formed at the nodes. They are important in vegetative reproduction. Fig. 20 AA AMOAH 25 3. Specialized functions by certain metamorphosed aerial organs. E.g. Climbing stems: Examples are pole bean, yam and many vines in tropical rain forest. Some of these species merely leans upon other plants, fences and rocks. Others climb by coiling around a support or by special modification such as tendrils. NB: Branching: In many plants especially dicots, the lateral buds develop into lateral shoots or branches, but in cereals and other grasses this growth in plant is modified. In these plants, the main axis of the shoot usually produces branches only from the basal nodes i.e. those close to or just beneath the surface of the soil. Lateral buds in the axils of the leaves at the base of the main shoot give rise to secondary aerial branches. The secondary shoots intend may produce branches of a third order from basal nodes, this process may continue until a considerable number of branches are formed. This method of branching i.e. from basal nodes is called tillering and the resulting branches are known as tillers (e.g. rice plant). The number of tillers formed depends on heredity or variety, closeness of planting or spacing and environmental factors such as moisture availability. Tillering and branching are of great agronomic importance and the agriculturist depends on this reaction to multiply number of heads, pods or ears per plant. AA AMOAH 26