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1 INTRODUCTION What is botany? Botany is the branch of biology concerned with the scientific study of plants. Traditionally, botanists studied all organisms that were not generally regarded as animal. However, advances in our knowledge about the myriad forms of life, especially microbes (viruses and bacteria), have led to spinning off from Botany the specialized field called Microbiology. Still, the microbes are usually covered in introductory Botany courses, although their status as neither animal nor plant is firmly established. • Importance of plants • Photosynthesis • Produces food. Photosynthesis feeds the plant but also feeds heterotrophic organisms (like us) that eat the plant. About 200,000,000,000 tons of carbon per year are “fixed” by plants. That’s 6,000 tons per second. • Produces oxygen. Plants also produce oxygen needed by aerobic species (like us). • Plant products people use • Food • Spices • Wood • Textiles • Drugs 2 • Fossil fuels • Aesthetics • Research. Many basic principles of biology have been discovered through study of plants. • Plants are more like animals than they are different • There bodies are composed of cells • They carry out most of the same biochemical reactions. such as glycolysis and the Krebs cycle, protein synthesis, etc., etc. • Their genes are composed of DNA • Their cells divide by mitosis and meiosis • Most reproduce by sexual reproduction • They evolve by natural selection • Plants are simple in structure • They are much simpler than higher animals. Plants have only 7 basic types of tissues and only three main organs. • Plants are chemically complex • Chemically, plants are more complex than animals. Plants can produce all of the thousands of organic compounds needed for life from water, carbon dioxide, and a few minerals. • The branches of botany Some branches based on general properties of plants • Taxonomy • Morphology • Anatomy 3 • Physiology • Ecology • Genetics • Cell biology Some branches based on kind of plant • Phycology-algae • Mycology-fungi • Bryology-mosses • Pterology-ferns What is a plant? • Two-kingdom system. • Originally, all living organisms were classified in two kingdoms • Plant Kingdom - How do we define “plant?” • Autotrophic • Nonmotile • Cells have cell walls • Animal Kingdom - How do we define “animal?” • Heterotrophic • Motile • Cells lack walls • Problems. • As new organisms were described (especially microscopic ones) it was recognized that many species did not easily fit into “plant” or “animal.” 4 • Examples of “problem” organisms • Euglena. Motile (animal?), no cell wall (animal?), but photosynthetic (plant?) • Slime molds. Motile (animal?), heterotrophic (animal?), spores have walls (plant?) • Mushrooms. Cell walls (plant?), nonmotile (plant?), but heterotrophic (animal?) • Five-kingdom system. • This is one solution to the “problem” organisms • Kingdom Monera. Cells are prokaryotic. Includes bacteria and cyanobacteria. • Kingdom Protista. A complex group of “simple” eukaryotic organisms. The plant-like protists are the algae. They are photosynthetic but much simpler than higher plants. • Kingdom Myceteae (Kingdom Fungi). These are eukaryotic species with cell walls but lacking photosynthesis. Includes yeasts, molds, mushrooms. • Kingdom Plantae. The so-called “land plants.” Most of these fit the usual definition of plant. They are photosynthetic like algae but more complex in structure. • Kingdom Animalia. Higher animals (we will let the zoology class cover these). by Dr. John Tiftickjian [[email protected]] Sciences). 5 . Delta State University (Biological Plant cell structure 6 Plants are unique among the eukaryotes, organisms whose cells have membrane-enclosed nuclei and organelles, because they can manufacture their own food. Chlorophyll, which gives plants their green color, enables them to use sunlight to convert water and carbon dioxide into sugars and carbohydrates, chemicals the cell uses for fuel. Cell wall A thick, rigid membrane that surrounds a plant cell. This layer of cellulose fiber gives the cell most of its support and structure. The cell wall also bonds with other cell walls to form the structure of the plant. Cell membrane The thin layer of protein and fat that surrounds the cell, but is inside the cell wall. The cell membrane is semipermeable, allowing some substances to pass into the cell and blocking others. Plasmodesmata Plasmodesmata are small tubes that connect plant cells to each other, providing living bridges between cells. Chloroplasts The most important characteristic of plants is their ability to photosynthesize, in effect, to make their own food by converting light energy into chemical energy. This process is carried out in specialized organelles called chloroplasts. Endoplasmic reticulum The endoplasmic reticulum is a network of sacs that manufactures, processes, and transports chemical compounds for use inside and outside of the cell.. In plants, the endoplasmic reticulum also connects between cells via the plasmodesmata. 7 Rough endoplasmic reticulum - (rough ER) a vast system of interconnected, membranous, infolded and convoluted sacks that are located in the cell's cytoplasm (the ER is continuous with the outer nuclear membrane). Rough ER is covered with ribosomes that give it a rough appearance. Rough ER transport materials through the cell and produces proteins in sacks called cisternae (which are sent to the Golgi body, or inserted into the cell membrane). Smooth endoplasmic reticulum - (smooth ER) a vast system of interconnected, membranous, infolded and convoluted tubes that are located in the cell's cytoplasm (the ER is continuous with the outer nuclear membrane). The space within the ER is called the ER lumen. Smooth ER transport materials through the cell. It contains enzymes and produces and digests lipids (fats) and membrane proteins; smooth ER buds off from rough ER, moving the newly-made proteins and lipids to the Golgi body and membranes. Golgi body Also called the golgi apparatus or golgi complex) a flattened, layered, sac-like organelle that looks like a stack of pancakes and is located near the nucleus. The golgi body packages proteins and carbohydrates into membrane-bound vesicles for "export" from the cell. Mitochondria Spherical to rod-shaped organelles with a double membrane. The inner membrane is infolded many times, forming a series of projections (called cristae). The mitochondrion converts the energy stored in glucose into ATP (adenosine triphosphate) for the cell. Nucleus 8 Spherical body containing many organelles, including the nucleolus and surrounded by nuclear membrane. The nucleus is a highly specialized organelle that serves as the information processing and administrative center of the cell. This organelle has two major functions: it stores the cell's hereditary material, or DNA, and it coordinates the cell's activities, which include growth, intermediary metabolism, protein synthesis, and reproduction (cell division). Nucleolus An organelle within the nucleus - it is where ribosomal RNA is produced. Nuclear membrane The membrane that surrounds the nucleus. Vacuole Each plant cell has a large, single vacuole that stores compounds, helps in plant growth, and plays an important structural role for the plant. Ribosomes Small organelles composed of RNA-rich cytoplasmic granules that are sites of protein synthesis. Cytoplasm The jellylike material outside the cell nucleus in which the organelles are located. Microtubules These straight, hollow cylinders are found throughout the cytoplasm of all eukaryotic cells (prokaryotes don't have them) and carry out a variety of functions, ranging from transport 9 to structural support. Plant Tissues Plants are composed of three major organ groups: roots, stems and leaves. As we know from other areas of biology, these organs are comprised of tissues working together for a common goal (function). In turn, tissues are made of a number of cells which are made of elements and atoms on the most fundamental level. In this section, we will look at the various types of plant tissue and their place and purpose within a plant. It is important to realize that there may be slight variations and modifications to the basic tissue types in special plants. Plant tissues are characterized and classified according to their structure and function. The organs that they form will be organized into patterns within a plant which will aid in further classifying the plant. A good example of this is the three basic tissue patterns found in roots and stems which serve to delineate between woody dicot, herbaceous dicot and monocot plants. We will look at these classifications later on in the tutorial. 10 Classification of Plant tissues Plant tissues Meristematic tissues Primary Permanent tissues Secondary Simple Complex Dermal Parenchyma Collenchyma Sclerenchyma 11 Vascular A- Meristematic Tissues Tissues where cells are constantly dividing are called meristems or meristematic tissues. These regions produce new cells. These new cells are generally small, six-sided boxlike structures with a number of tiny vacuoles and a large nucleus, by comparison. Sometimes there are no vacuoles at all. As the cells mature the vacuoles will grow to many different shapes and sizes, depending on the needs of the cell. It is possible that the vacuole may fill 95% or more of the cell’s total volume. There are 3 types of meristem Apical meristem Lateral meristem Intercalary meristem -- Apical meristems are located at or near the tips of roots and shoots. As new cells form in the meristems, the roots and shoots will increase in length. This vertical growth is also known as primary growth. A good example would be the growth of a tree in height. Each apical meristem will produce embryo leaves and buds as well as three types of primary meristems: protoderm, ground meristems, and procambium. These primary meristems will produce the cells that will form the primary tissues. 12 Root apex 13 Soot apex 14 -- Lateral meristems account for secondary growth in plants. Secondary growth is generally horizontal growth. A good example would be the growth of a tree trunk in girth. There are two types of lateral meristems to be aware of in the study of plants. 1- The vascular cambium, the first type of lateral meristem, is sometimes just called the cambium. The cambium is a thin, branching cylinder that, except for the tips where the apical meristems are located, runs the length of the roots and stems of most perennial plants and many herbaceous annuals. The cambium is responsible for the production of cells and tissues that increase the thickness, or girth, of the plant. 2- The cork cambium, the second type of lateral meristem, is much like the vascular cambium in that it is also a thin cylinder that runs the length of roots and stems. The difference is that it is only found in woody plants, as it will produce the outer bark. Both the vascular cambium and the cork cambium, if present, will begin to produce cells and tissues only after the primary tissues produced by the apical meristems have begun to mature. -- Intercalary meristems are found in grasses and related plants that do not have a vascular cambium or a cork cambium, as they do not increase in girth. These plants do have apical meristems and in areas of leaf attachment, called nodes, they have the third type of meristematic tissue. This meristem will also actively produce new cells and is responsibly for increases in length. The intercalary meristem is responsible for the regrowth of cut grass. B- Permanent tissues 15 Tissues in plants that do not actively produce new cells. These tissues are called nonmeristematic tissues (permanent tissues). Nonmeristematic tissues are made of cells that are produced by the meristems and are formed to various shapes and sizes depending on their intended function in the plant. Sometimes the tissues are composed of the same type of cells throughout, or sometimes they are mixed. There are simple tissues and complex tissues. 1- Simple tissues There are three basic types, named for the type of cell that makes up their composition. Parenchyma (cells with thin primary walls that retain their protoplasm) Parenchyma is the most common and versatile ground tissue. It forms, for example, the cortex and pith of stems, the cortex of roots, the mesophyll of leaves, the pulp of fruits, and the endosperm of seeds. Parenchyma cells are living cells and may remain meristematic at maturity, meaning that they are capable of cell division. They have thin but flexible cellulose cell walls, and are generally polygonal when close-packed, but approximately spherical when isolated from their neighbours. They have large central vacuoles, which allows the cells to store and regulate ions, waste products and water. Parenchyma cells have a variety of functions: In leaves, they form the mesophyll and are responsible for photosynthesis and the exchange of gases[1], parenchyma cells in 16 the mesophyll of leaves are a specialized parenchymatous tissue known as chlorenchyma (parenchyma with chloroplasts). Storage of starch, protein, fats and oils and water in roots, tubers (e.g. potato), seed endosperm (e.g. cereals) and cotyledons (e.g. pulses and groundnut) Secretion (e.g. hydathodes, nectaries and cells lining the inside of resin ducts) Wound repair and the potential for renewed meristematic activity Other specialized functions such as aeration (aerenchyma) and support. 17 Collenchyma (cells with thick primary walls that retain their protoplasm) cells form collenchyma tissue. These cells have a living protoplasm, like parenchyma cells, and may also stay alive for a long period of time. Their main distinguishing difference from parenchyma cells is the increased thickness of their walls. In cross section, the walls looks uneven. Collenchyma cells are found just beneath the epidermis and generally they are elongated and their walls are pliable in addition to being strong. As a plant grows these cells and the tissues they form, provide flexible 18 support for organs such as leaves and flower parts. Collenchyma tissue have the following features. Collenchyma tissue provide flexible support while alive. Collenchyma cells are supporting cells. Their primary walls are characteristically thick at the corners of the cells. Collenchyma cells are generally elongated. In collenchyma cells, the primary wall thickens, but no secondary wall forms. Collenchyma provides support to leaf petioles, nonwoody stems, and growing organs. Tissue made of collenchyma cells is flexible, permitting stems and petioles to sway in the wind without snapping. There are three principal types of collenchyma: Angular collenchyma (thickened at intercellular contact points) Tangential collenchyma (cells arranged into ordered rows and thickened at the tangential face of the cell wall) Lacunar collenchyma (have intercellular space and thickening proximal to the intercellular space) Cross section of collenchyma cells 19 Diagram of Collenchyma Tissue 20 21 Sclerenchyma (cells with lignified secondary walls that have lost their protoplasm at maturity, i.e. are 'dead') Sclerenchyma is a supporting tissue in plants. Two groups of sclerenchyma cells exist: fibres and sclereids. Their walls consist of cellulose, hemicellulose and lignin. Sclerenchyma cells are the principal supporting cells in plant tissues that have ceased elongation. Sclerenchyma fibres are of great economical importance, since they constitute the source material for many fabrics Unlike the collenchyma, mature sclerenchyma is composed of dead cells with extremely thick cell walls (secondary walls) that make up to 90% of 22 the whole cell volume. It is the hard, thick walls that make sclerenchyma cells important strengthening and supporting elements in plant parts that have ceased elongation. The difference between fibres and sclereids is not always clear. Transitions do exist, sometimes even within one and the same plant. 1- Fibers Fibers are sometimes found in association with a wide variety of tissues in roots, stems, leaves and fruits. Usually fiber cells are much longer than they are wide and have a very tiny cavity in the center of the cell. Currently, fibers from over 40 different plant families are used in the manufacture of textiles, ropes, string. Fibres usually originate from meristematic tissues. Cambium and procambium are their main centers of production. They are usually associated with the xylem and phloem of the vascular bundles. The fibres of the xylem are always lignified, while those of the phloem are cellulosic. Reliable evidence for the fibre cells' evolutionary origin from tracheids exists. During evolution the strength of the tracheid cell walls was enhanced, the ability to conduct water was lost and the size of the pits reduced. Fibres that do not belong to the xylem are bast (outside the ring of cambium) and such fibres that are arranged in characteristic patterns at different sites of the shoot. 23 Cross section of sclerenchyma fibers 2- Sclereids Sclereids are sclerenchyma cells that are randomly distributed throughout other tissues. Sometimes they are grouped within other tissues in specific zones or regions. They are generally as long as they are wide. An example, would be the gritty texture in some types of pears. The grittiness is due to groups of sclereid cells. Sclereids are sometimes called stone cells. Fresh mount of a sclereid Moore, Randy; Clark, W. Dennis; and Vodopich, Darrell S. (1998). Botany (3rd ed.). 24 2- Epidermal tissue system The epidermal tissue includes several differentiated cell types: epidermal cells, guard cells, and epidermal hairs (trichomes). The epidermal cells are the most numerous, largest, and least specialized. These are typically more elongated in the leaves of monocots than in those of dicots. --- Epidermis The epidermis is a single-layered group of cells that covers plants' leaves, flowers, roots and stems. It forms a boundary between the plant and the external world. The epidermis serves several functions, it protects against water loss, regulates gas exchange, secretes metabolic compounds, and (especially in roots) absorbs water and mineral nutrients. The epidermis of most leaves shows dorsoventral anatomy: the upper (adaxial) and lower (abaxial) surfaces have somewhat different construction and may serve different functions. The walls of the epidermal cells of the above ground parts of plants contain cutin, and are covered with a cuticle. The cuticle reduces water loss to the atmosphere, it is sometimes covered with wax in smooth sheets or long filaments. Woody stems and some other stem structures produce a secondary covering called the periderm that replaces the epidermis as the protective covering. 25 --- Trichomes Trichomes or hairs grow out from the epidermis in many species. In root epidermis, epidermal hairs, termed root hairs are common and are specialized for absorption of water and mineral nutrients. Plant hairs may be unicellular or multicellular , branched or unbranched. Multicellular hairs may have one or several layers of cells. 26 Ascensão, L., N. Marques and M.S. Pais. 1995. Glandular trichomes on vegetative and reproductive organs of Leonotis leonurus (Lamiaceae). Annals of Botany 75: 619-626. --- Stomata An important feature of leaf epidermis is the presence of stomata which occur either on both sides of the leaf or only on one side of the leaf. When stomata occur on both sides, the leaf is known as amphistomatic, 27 when they are confined to the upper side, the leaf is known as epistomatic and when to the lower side, the leaf is called hypostomatic. Each stomata consists of an opening (pore) bounded by two specialized, usually kidney shaped, epidermal cells known as guard cells. The guard cells have unevenly thickened walls. The inner wall facing the aperture is highly thickened while the one away from the aperture is thin and extensible. The guard cells are also covered with cuticle which extends to the inner wall forming the boundary of the pore and the sub-stomatal chamber. Guard cells are surrounded by a variable number of epidermal cells which are called subsidiary or accessory cells. These cells may be morphologically similar to the other epidermal cells or very different from them. Raven, Peter H.; Evert, Ray F.; Curtis, Helena (1981), Biology of plants, New York, N.Y.: Worth Publishers, pp. 427–28 28 29 3- Vascular tissues Vascular tissue is a complex conducting tissue, formed of more than one cell type, found in vascular plants. The primary components of vascular tissue are the xylem and phloem. These two tissues transport fluid and nutrients internally. There are also two meristems associated with vascular tissue: the vascular cambium and the cork cambium. All the vascular tissues within a particular plant together constitute the vascular tissue system of that plant. 30 Xylem It is a complex permanent tissue, which is specialized for the conduction of water and mineral substances in the plant body. Xylem is a heterogenous tissue made up of four different types of cellular elements. They are: Xylem tracheids Xylem tracheae Xylem fibers and Xylem parenchyma 1- Xylem Tracheids They are found abundantly in pteridophytes, gymnosperms and primitive angiosperms. In these groups of plants, the tracheids represent the most active water conducting elements. In advanced angiosperms, the tracheids are found restricted to leaf margin and leaf tip. The tracheids are elongated, dead cells, with tapering ends. They are characterised by the presence of a thick cell wall consisting of primary wall and a secondary wall. The primary wall is composed of cellulose where as the secondary wall is made up of lignin. There is a spacious lumen that extends throughout the length of the tracheid. In some cases, due to the deposition of lignin, the primary wall develops numerous concave depressions called pits. When pits are present, the tracheid is described as pitted and when pits are absent, it is described as simple. 31 Tracheids 32 2- Xylem Tracheae They are commonly known as xylem vessels. They are the most active water conducting elements in all higher angiosperms. The tracheae are found arranged parallel to each other, extending from one end of the plant body to another. The tracheae are long cylindrical dead cells. They are characterised by a thick cell wall consisting of a primary wall and a secondary wall. The primary wall is made up of cellulose where as the secondary wall is made up of lignin. There is a spacious lumen that extends throughout the length of the trachea. The deposition of lignin in the secondary wall is not always uniform. As a result, the xylem vessels exhibit different types of secondary thickenings. On this basis, xylem vessels can be distinguished into five types: Annular vessels in which the secondary thickening is in the form of rings placed more or less at equal distance from each other. Spiral vessels in which the secondary thickenings are present in the form of a helix or coil. Scalariform vessels in which the secondary thickenings appear in the form of cross bands resembling the steps of a ladder. Reticulate vessels in which the secondary thickenings are irregular and appear in the form of a network. Pitted vessels in which the secondary thickenings result in the 33 B-G: trachea in LS showing different types of thickenings 34 3- Xylem Fibres They are represented by the dead sclerenchyma fibers that are found in between the vessels and the tracheids. They are meant for providing mechanical support to the essential elements. 4- Xylem Parenchyma This is the only living component in the xylem tissue. It is represented by groups of parenchyma cells that are found in between the vessels and the fibers. They are meant for storage of reserve food. Types of Xylem Xylem can be distinguished into two types namely Primary xylem and Secondary xylem Primary Xylem Primary xylem is the xylem that is formed during normal growth. It is a derivative of primary meristem. It occurs in both monocots and dicots. In the primary xylem, two types of xylem vessels can be distinguished, namely protoxylem and metaxylem. 35 Secondary Xylem Secondary xylem is the xylem that is formed during secondary growth. It is derivative of secondary meristem. It is a characteristic feature of only dicots. Secondary xylem is commonly known as wood. It is of commercial importance since it is extensively used in the manufacturing of doors, windows and furniture. Wilson, K. & D.J.B. White (1986). The Anatomy of Wood: its Diversity and variability. Stobart & Son Ltd, London Peter A. Raven, Ray F. Evert, Susan E. Eichhorn (1999). Biology of Plants. W.H. Freeman and Company. pp. 576-577. Kenrick, Paul; Crane, Peter R. (1997). The Origin and Early Diversification of Land Plants: A Cladistic Study. Washington, D. C.: Smithsonian Institution Press. ISBN 1-56098-730-8 Campbell, Neil A.; Jane B. Reece (2002). Biology (6th ed.). Benjamin Cummings. ISBN 978-0805366242. Melvin T. Tyree; Martin H. Zimmermann (2003). Xylem Structure and the Ascent of Sap (2nd ed.). Springer. ISBN 3-540-43354-6. recent update of the classic book on xylem transport by the late Martin Zimmermann. Phleom Phloem is a complex permanent tissue, which is specialized for the conduction of food and other organic substances. Phloem is also a 36 heterogenous tissue, made up of four different types of cellular elements, namely, Sieve tubes Companion cells Phloem parenchyma and Phloem fibres Sieve Tubes They represent the most active food conducting elements in the phloem tissue. The sieve tubes are found arranged parallel to one another from one end of the plant body to another. Each sieve tube is formed by a series of hollow, cylindrical cells called sieve tube cells arranged one above the other. The sieve cells are separated from each other by horizontal perforated plates called sieve plates. The sieve cells communicate with each other through the sieve plates. Each sieve cell has a thin cell wall, which is composed of only cellulose. The cell has a central mass of dense cytoplasm. The granular cytoplasm forms a number of projections called cytoplasmic strands extending towards the sieve plate. The nucleus is absent. Companion Cells They are more or less spindle shaped cells associated with the sieve tube cells. Each companion cell is found attached to any one lateral surface of a sieve cell. The companion cell and the neighbouring sieve cell together represent a pair of sister cells. The companion cell has a granular cytoplasm, prominent nucleus and one or two small vacuoles. The 37 nucleus of the companion cell is said to be capable of exerting its influence on the adjacent sieve cells. Phloem Structure Phloem Parenchyma Phloem parenchyma is represented by a group of living parenchyma cells that are found in-between the sieve tubes. They are meant only for storage of organic food Phloem Fibres Phloem fibres are represented by the dead sclerenchyma fibres that are found in between the sieve tubes. They are meant only for providing mechanical support. Types of Phloem 38 Phloem can be distinguished into two types namely Primary phloem Secondary phloem Primary Phloem Primary phloem is the phloem that is formed during normal growth in the plant body. It is a derivative of primary meristem. It is found in both monocots and dicots. The primary phloem is further composed of protophloem and metaphloem. The sieve tubes and the companion cells, which appear earlier during normal growth, represent protophloem, while metaphloem is represented by the sieve tubes and companion cells that appear later. However, there is no significant morphological difference between protophloem and metaphloem. Secondary Phloem Secondary phloem is the phloem that is formed during secondary growth. It is a derivative of secondary meristem. Secondary phloem is characteristic feature of only dicots. It is also known as bast. It is also of commercial importance since it yields bast fibers. 39 The following table summarise the characteristics of and difference between xylem and phloem: Characteristics Xylem Phloem Dead, complex permanent Living, complex tissue permanent tissue Xylem tracheids and Sieve tubes and tracheae companion cells Associated Xylem fibres and xylem Phloem fibres and elements parenchyma phloem parenchyma Xylem tracheids Only Phloem fibres Definition Essential elements Non-living components Living components Function Sieve tubes companion Only xylem parenchyma cells and phloem parenchyma Conduction of water 40 Conduction of food and other organic substances Types 1. Primary xylem 1.Primary phloem (a) protoxylem (a) protopholoem (b) metaxylem (b) metaphloem 2. Secondary xylem (or 2. Secondary pholem (or wood) bast) 41 Secondary growth The normal process of growth that occurs in every plant body is known as primary growth. It is the result of the activity of primary meristem. The process of primary growth results in the formation of primary permanent tissues such as primary xylem, primary phloem and primary cortex. However in the dicot plants, there is a process of growth that begins after a known period of primary growth. Such a growth is known as secondary growth. It is the result of the activity of secondary meristem. It results in the formation of secondary permanent tissues such as secondary xylem, secondary phloem and secondary cortex. As a result, secondary growth brings about an increase in the girth of the plant body. Some Important Definitions: Primary tissues: Tissues generated from the growth of an apical meristem. Cambium: A lateral meristem constituting a sheet of cells. Growth of these cells increases the girdth of the plant organ involved. Secondary tissues: Tissues generated from the growth of a cambium. Vascular Cambium: A cambium that gives rise to secondary xylem to the inside, and to secondary phloem to the outside. Two kinds of meristematic cells, called initials, are recognizable in the cambium: fusiform and ray initials. The fusiform initials are elongated vertically in the stem and have tapering ends. They divide to produce the conducting cells of both the xylem and the phloem (xylem toward the inside of the stem, phloem toward the outside). 42 Considerably more xylem cells than phloem cells always are produced. The ray initials are smaller, more cuboidal and produce parenchyma in rows radiating out from the center of the stem. The bands of parenchyma, called rays (vascular rays), conduct water and dissolved materials laterally in the stem. Periderm: A structure that consists of a cork cambium (phellogen), with cork tissue (phellem) to the outside, and in some cases a layer of cells derived from and to the inside of the cork cambium called phelloderm. Functions to limit dehydration and block pathogens after the epidermis is disrupted by the onset of secondary growth. Cork: (phellem) you need know only the term "cork": Tissue dead at maturity generated from a cork cambium. The cell walls of the tissue are impregnated with suberin. This water-proofs the tissue. Cork Cambium: A cambial layer that functions to produce cork, and in some cases, phelloderm. In roots is derived initially from pericyle. In stems from the cortex. Unlike the vascuar cambium these cambial layers do not persist for the duration of the life of the plant organ. Over time one cork cambium will be supplanted by another generated from parenchyma cells further inside. Phelloderm: In some periderms a layer of living secondary tissue is generated by the cork cambium to the inside. We will not consider thie phelloderm in the following exercise. 43 44 Characteristics of Diagram of young dicot stem 1- Ground tissue consists of cortex and pith 2- Cortex is characterized into collenchyma and parenchyma 3- The vascular bundles arranged in ring 4- The vascular bundle is open 5- The phloem is irregular 6- Metaxylem and protoxylem arranged in rows 7- The vascular bundle is lateral 45 Changes accompanied to secondary growth in dicot stem 46 After significant activity in the vascular cambium, a stem exhibiting secondary growth might look like the previous diagram. The primary xylem is in the center of the stem, while the primary phloem is pushed outward by the new cells that arise from the vascular cambium. Eventually, the primary phloem is crushed into the cortex. The secondary xylem differentiates from the cells that divide off the vascular cambium towards the inside of the stem, while the secondary phloem differentiates from the cells that divide towards the outside of the stem. Thompson, N.P. and Heimsch, C. 1964. Stem anatomy and aspects of development in tomato. American Journal of Botany 51: 7-19. Esau, K. and Cheadle, V.I. 1969. Secondary growth in bougainvillea. Annals of Botany 33: 807-819. Ewers, F.W. 1982. Secondary growth in needle leaves of Pinus longaeva (bristlecone pine) and other conifers: Quantitative data. American Journal of Botany 69: 1552-1559. 47 Secondary growth in roots Secondary growth in roots begins with the formation of the vascular cambium. This concentric, undifferentiated cell layer originates from the pericycle and from the procambium, which is located between the primary xylem and primary phloem tissues. The vascular cambium continues to divide and differentiate to produce secondary vascular tissue toward the center of the root and secondary phloem tissue toward the outside of the root. The pericycle also gives rise to the cork cambium, which produces the periderm to the outside of the root. Primary and secondary xylem in alternate position. Primary and secondary phleom next to each other. 48 Photosynthesis Photosynthesis is the process of converting light energy to chemical energy and storing it in the bonds of sugar. This process occurs in plants and some algae. Plants need only light energy, CO 2, and H2O to make sugar. The process of photosynthesis takes place in the chloroplasts, specifically using chlorophyll, the green pigment involved in photosynthesis. Photosynthesis takes place primarily in plant leaves, and little to none occurs in stems, etc. The parts of a typical leaf include the upper and lower epidermis, the mesophyll, the vascular bundle(s) (veins), and the stomates. The upper and lower epidermal cells do not have chloroplasts, thus photosynthesis does not occur there. They serve primarily as protection for the rest of the leaf. The stomates are holes which occur primarily in the lower epidermis and are for air exchange: they let CO 2 in and O2 out. The vascular bundles or veins in a leaf are part of the plant's transportation system, moving water and nutrients around the plant as needed. The mesophyll cells have chloroplasts and this is where photosynthesis occurs. 49 50 Photosynthesis occurs in chloroplasts Chloroplasts Chloroplasts are probably the most important among the plastids since they are directly involved in photosynthesis. They are usually situated near the surface of the cell and occur in those parts that receive sufficient light, e.g. the palisade cells of leaves. The green colour of chloroplasts is caused by the green pigment chlorophyll. Structure Chloroplasts are usually disc-shaped and surrounded by a double membrane. Inside the inner membrane there is a watery protein-rich ground substance or stroma in which is embedded a continuous membrane system, the granal network. This network forms a threedimensional arrangement of membrane-bound vesicles called thylakoids. The thylakoids usually lie in stacks called grana and contain the photosynthetic pigments - green chlorophyll a and b and the yellow to red carotenoids. The grana are interconnected by tubular membranes called the intergranal frets or lamellae. Functions o chloroplasts are the sites for photosynthesis; o they contain enzymes and co-enzymes necessary for the process of photosynthesis. 51 52 Stages of photosynthesis Light reactions Dark reactions (Calvin cycle) Light reactions The light-dependent reactions require light. These reactions occur in the thylakoid membrane. They produce Energy storage molecules ATP and NADPH, which are needed to produce glucose in the light-independent reactions Oxygen gas is made as a waste product. 2 H2O + 2 NADP+ + 3 ADP + 3 Pi + light → 2 NADPH + 2 H+ + 3 ATP + O2 12 H2O + Energy 6 O2 + 24 H+ + 24e- 53 Dark reactions Dark reactions (light-independent) occur in the stroma. Carbon dioxide is “fixed” into the sugar glucose. ATP and NADPH molecules created during the light reactions power the production of this glucose. 3 CO2+ 9 ATP + 6 NADPH + 6 H+ → C3H6O3-phosphate + 9 ADP + 8 Pi + 6 NADP+ + 3 H2O 6 CO2 + 24 H+ + 24 e- ------> C6H12O6 + 6 H2O 2 Sets of Reactions 54 Factors affecting the rate of photosynthesis Different factors affect the rate of photosynthesis such as External (sulight, carbondioxide, temperature, water and oxygen) and internal (age of the leaf, number of stomata, chlorophyll etc). External Factors Affecting Photosynthesis: 1- Light Wavelength o Red and blue light are most effective for driving photosynthesis. Green (500nm) is least effective. 2. Light Intensity o Photosynthesis goes faster in more intense light until limited by some other factor. Once the reactions are going as fast as they can, more intense light has no effect. 3. Temperature o Up to a point, the rate of photosynthesis increases with temperature. Above the optimum temperature for photosynthetic enzyme function, photosynthesis is inhibited or shut down completely. 4. Humidity o If the humidity surrounding a plant is low, the stomata will close to reduce water loss through transpiration. Closed stomata limit gas exchange, and photosynthesis is slowed by a reduction in carbon dioxide availability. 5. Carbon Dioxide Concentration 55 o More carbon dioxide in the air allows more photosynthetic conversion into sugar, until limited by another factor. This is why some growers add carbon dioxide to the air in a growing environment. 6. Water o Compared to the amount of water needed to sustain a plant, the amount needed for photosynthesis is small. However, a dehydrated plant will suffer physical damage and will also close stomata, thus limiting photosynthesis. Internal factors of photosynthesis 1. Chlorophyll concentration --The concentration of chlorophyll affects the rate of reaction as they absorb the light --Lack or deficiency of chlorophyll results in chlorosis or yellowing of leaves --Lack of iron, magnesium, nitrogen and light affect the formation of chlorophyll 2. Age --Rate of photosynthesis increases with age of the leaf till maturation. Afterwards it begins to decrease. 3. Hormones --Cytokinin and gibberellins enhance the rate of photosynthesis while ABA decreases. 4. Carotenoids --Essential to prevent photo-oxidation which reduces photosynthesis 56 5. Translocation --The energy rich carbon compounds formed during photosynthesiscalled photosynthates or photoassimilates are transported to all the organs and tissues of the plant body. This long distance transport of photosynthates through the phloem is called as translocation. --The photosynthates provide energy to the non-photosynthetic tissues through respiration. In storage organs, photosynthates are stored in the form of starch or as other carbohydrates. Bidlack JE; Stern KR, Jansky S (2003). Introductory plant biology. New York: McGraw-Hill. Reece, J, Campbell, N (2005). Biology. San Francisco: Pearson, Benjamin Cummings. Govindjee Beatty JT,Gest H, Allen JF (2006). Discoveries in Photosynthesis. Advances in Photosynthesis and Respiration. 20. Berlin: Springer. Blankenship RE (2008). Molecular Mechanisms of Photosynthesis (2nd ed.). John Wiley & Sons Inc. Plant physiology, (4th edition ) M.Devlin, Francis H. Withan 1983. 57 Transpiration Transpiration is the evaporation of water into the atmosphere from the leaves and stems of plants. Plants absorb soilwater through their roots and this water can originate from deep in the soil. Plants pump the water up from the soil to deliver nutrients to their leaves. This pumping is driven by the evaporation of water through small pores called "stomates", which are found on the undersides of leaves. Transpiration accounts for approximately 10% of all evaporating water. Importance supply photosynthesis (1%-2% of the total) bring minerals from the roots for biosynthesis within the leaf cool the leaf 58 Environmental factors that affect the rate of transpiration 1. Light Plants transpire more rapidly in the light than in the dark. This is largely because light stimulates the opening of the stomata (mechanism). Light also speeds up transpiration by warming the leaf. 2. Temperature Plants transpire more rapidly at higher temperatures because water evaporates more rapidly as the temperature rises. At 30°C, a leaf may transpire three times as fast as it does at 20°C. 3. Humidity The rate of diffusion of any substance increases as the difference in concentration of the substances in the two regions increases.When the 59 surrounding air is dry, diffusion of water out of the leaf goes on more rapidly. 4. Wind When there is no breeze, the air surrounding a leaf becomes increasingly humid thus reducing the rate of transpiration. When a breeze is present, the humid air is carried away and replaced by drier air. 5. Soil water A plant cannot continue to transpire rapidly if its water loss is not made up by replacement from the soil. When absorption of water by the roots fails to keep up with the rate of transpiration, loss of turgor occurs, and the stomata close. This immediately reduces the rate of transpiration (as well as of photosynthesis). If the loss of turgor extends to the rest of the leaf and stem, the plant wilts. Benjamin Cummins (2007), Biological Science (3 ed.), Freeman, Scott, p. 215 Debbie Swarthout and C.Michael Hogan. 2010. Stomata. Encyclopedia of Earth 60 Absorption Absorption of Water by Plants Plants absorb water through the entire surface - roots, stems and leaves. However, mainly the water is absorbed by roots. The area of young roots where most absorption takes place is the root hair zone. The root hairs are delicate structures which get continuously replaced by new ones at an average rate of 100 millions per day. The root hairs lack cuticle and provide a large surface area. They are extensions of the epidermal cells. They have sticky walls by which they adhere tightly to soil particles. As the root hairs are extremely thin and large in number, they provide enormous surface area for absorption. They take in water from the intervening spaces mainly by osmosis. 61 62 Absorption of Water & Minerals in Plants The absorption of both water and minerals is important for the health, growth and fruit production of plants. Water and minerals are absorbed by the plant's roots, a process which is often aided by specialized fungi that form symbiotic relationships with the roots that are known as mycorrhizae. Roots The outermost cell layer of a root is known as the epidermis. It is these cells that absorb water from the environment as well as minerals that are dissolved in the water. Just past the growing tip of the root, some of the epidermal cells are elongated. These cells function to increase the surface area of the root and thereby improve its capacity to absorb water and minerals. Root hairs are easily torn off or desiccated by the sun when plants are transplanted, thereby temporarily reducing the capacity of the plant to absorb water and minerals. Mycorrhizae Where mycorrhizal relationships exist, the fungus puts out thin, thready hyphae into the surrounding soil. These absorptive hyphae increase the area of the root system and therefore its access to water and nutrients. The water and nutriets absorbed by the hyphae are exchanged for sugars produced by the plant. Mycorrhizal plants usually perform and survive better than those without this symbiotic relationship. Mycorrhizae have a vital function in the absorption of phosphorous and the prevention of iron and manganese deficiencies in alkaline soils. 63 Mineral Nutrition of Plants There are 14 essential mineral nutrients of plants, which are classified according to the amount required. Those for which plants have large requirements, or macronutrients, are nitrogen (N), potassium (K), phosphorus (P), calcium (Ca), sulfur (S) and magnesium (Mg). It is more common to see deficiencies of nitrogen, potassium and phosphorous than the other three minerals. Those minerals for which plants have smaller needs, or micronutrients are boron (B), chlorine (Cl), iron (Fe), copper (Cu), manganese (Mn), nickel (Ni), molybdenum (Mo) and zinc (Zn). The amount of minerals taken up by the plant depends both on the root's ability to absorb them and the concentration of nutrients around the root surface. Soil pH and Mineral Absorption Soil pH is a measure of the relative amounts of acid and alkaline ions in the soil. Equal amounts of each type of ion give a neutral soil with pH 7. The lower the pH, the more acid the soil, and the higher, the more alkaline. Phosphorous and various micronutrients are much more difficult for plants to absorb with a high-alkaline pH, and woody plants such as pin oak may show obvious symptoms of iron and manganese deficiency such as thin and pale leaves. Plants can generally tolerate a pH range of 5.5 to 8.3 if in a welldrained soil, but levels of 6.0.to 6.5 are optimal for most. 64 What is soil? Soil is a thin layer of material on the Earth's surface in which plants have their roots. It is made up of many things, such as weathered rock and decayed plant and animal matter. Soil is formed over a long period of time. Soil Formation takes place when many things interact, such as air, water, plant life, animal life, rocks, and chemicals. Soil formation Soil is formed from the weathering of rocks and minerals. The surface rocks break down into smaller pieces through a process of weathering and is then mixed with moss and organic matter. Over time this creates a thin layer of soil. Plants help the development of the soil. How? The plants attract animals, and when the animals die, their bodies decay. Decaying matter makes the soil thick and rich. This continues until the soil is fully formed. The soil then supports many different plants. Weathering: Weathering is the process of the breaking down rocks. There are two different types of weathering. Physical weathering and chemical weathering. In physical weathering it breaks down the rocks, but what it's made of stays the same. In chemical weathering it still breaks down the rocks, but 65 it may change what it's made of. For instance, a hard material may change to a soft material after chemical weathering. Stages in the Formation of Soil stage 1 stage 2 Soil composition Soils are a mixture of different things; rocks, minerals, and dead, decaying plants and animals. Soil can be very different from one location to another, but generally consists of organic and inorganic materials, water and air. The inorganic materials are the rocks that have been broken down into smaller pieces. The size of the pieces varies. It may appear as pebbles, gravel, or as small as particles of sand or clay. The organic material is decaying living matter. This could be plants or animals that have died and decay until they become part of the soil. The amount of water in the soil is closely linked with the climate and other characteristics of the region. The amount of water in the soil is one thing 66 that can affect the amount of air. Very wet soil like you would find in a wetland probably has very little air. The composition of the soil affects the plants and therefore the animals that can live there. Soil profile Soil Profile refers to the layers of soil; horizon A, B, and C. If you're wondering what horizon A is, here's your answer: horizon A refers to the upper layer of soil, nearest the surface. It is commonly known as topsoil. In the woods or other areas that have not been plowed or tilled, this layer would probably include organic litter, such as fallen leaves and twigs . The litter helps prevent erosion, holds moisture, and decays to form a very rich soil known as humus. Horizon A provides plants with nutrients they need for a great life. Soil conservation Soil erosion, caused by wind and rain, can change land by wearing down mountains, creating valleys, making rivers appear and disappear. It is a slow and gradual process that takes thousands, even millions of years. But erosion may be speeded up greatly by human activities such as farming and mining. Soil develops very slowly over a long period of time but can be lost too quickly. The clearing of land for farming, residential, and commercial use can quickly destroy soil. It speeds up the process of erosion by leaving soil exposed and also prevents development of new soil by removing the plants and animals that help build humus. Today's farmers try to farm in a way that reduces the amount of erosion and soil loss. They may plant cover crops or use a no-till method of 67 farming. Soil is an important resource that we all must protect. Without soil there is no life. Search Engine: http://www.yahooligans.com Soils, National Science Resource Center, 1996 Soil Types Sand, silt, and clay are the basic types of soil. Most soils are made up of a combination of the three. The texture of the soil, how it looks and feels, depends upon the amount of each one in that particular soil. The type of soil varies from place to place on our planet and can even vary from one place to another in your own backyard. 1- Sandy Soil Sand is the largest particle in the soil. The individual grains are large sized, thus added for increased aeration and easy draining. Nevertheless, sandy soil alone is not good for planting purpose, as it does not retain water, nutrient and fertilizers. 2- Silty Soil Silt is a soil particle whose size is between sand and clay. Silt feels smooth and powdery. When wet it feels smooth but not sticky. Similar to clay, silty soil turns sticky after saturation with water. 3- Clay Soil 68 Clay is the smallest of particles. Clay is smooth when dry and sticky when wet. Soils high in clay content are called heavy soils, sticky and the rate of saturation is high. . Clay also can hold a lot of nutrients, but doesn't let air and water through it well. This type of soil can be used after mixing with peat and sand particles. 4- Loamy Soil Loam is one of the ideal soil types for plant growing purposes. Generally, loam soil is fertile (unlike sandy soil) and has no water drainage problems like clayey soil and silty soil. In short, it is fertile and well-drained soil, excellent for cultivation. 5- Peat Soil Peat soil is loaded with organic materials (decaying remains of plants and animals). In comparison to other soil types, peat soil has the highest organic matter. Considering this, it is understandable that this is an acidic soil type (low pH range). Peat soil with moderately high pH and good water draining ability is good for plantation. 6- Shalky Soil In contrast to peat soil, the chalky soil type is alkaline in nature (high pH range) and prone to dryness. Also known as basic soil, it holds very less moisture, enabling water to drain off very easily. Despite the fact that chalky soil contains essential plant nutrients, they are not available to plants due to increased alkalinity. Ningthoujam Sandhyarani By Published: 15/10/2010 69