Anatomy, Embryology And Elementary Morphogenesis

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ANATOMY, EMBRYOLOGY AND ELEMENTARY MORPHOGENESISBSCBO-202Board of StudiesLate Prof. S. C. TewariDepartment of BotanyHNB Garhwal University,SrinagarProf. Uma PalniDepartment of BotanyRetired, DSB Campus,Kumoun University, NainitalDr. R.S. RawalScientist, GB Pant National Institute ofHimalayan Environment & SustainableDevelopment, AlmoraDr. H.C. JoshiDepartment of Environmental ScienceSchool of SciencesUttarakhand Open University,HaldwaniDr. Pooja JuyalDepartment of BotanySchool of SciencesUttarakhand Open University, HaldwaniProgramme CoordinatorDr. Pooja JuyalDepartment of BotanySchool of SciencesUttarakhand Open UniversityHaldwani, NainitalUnit Written By:Unit No.1. Dr. Prem PrakashAssistant Professor,Department of Botany,Govt. PG College Dwarahat1, 2, 3 & 42. Dr. Sushma TamtaAssistant Professor,Department of Botany,DSB Campus, Kumaun University, NainitalUTTARAKHAND OPEN UNIVERSITY5, 6 & 7Page 1

ANATOMY, EMBRYOLOGY AND ELEMENTARY MORPHOGENESIS3. Dr. Nishesh SharmaAssistant Prof., Department of Biotechnology,Chinmaya Degree College,BHEL, Haridwar4. Dr. Ritu V. SinghalAssistant Prof., Department of Botany,Chinmaya Degree College,BHEL, HaridwarBSCBO-2028 & 109Course EditorLate Prof. Y.P.S PangteyRetired Professor, Department of BotanyDSB Campus, Kumaun UniversityNainitalTitleISBN No.CopyrightEdition::::Anatomy, Embryology and Elementary MorphogenesisUttarakhand Open UniversityPublished By: Uttarakhand Open University, Haldwani, Nainital-263139UTTARAKHAND OPEN UNIVERSITYPage 2


ANATOMY, EMBRYOLOGY AND ELEMENTARY MORPHOGENESISBSCBO-202CONTENTSBLOCK-1GENERAL ANATOMYPAGE NO.Unit-1-Tools and Techniques in Plant Anatomy6-22Unit-2-Types of Tissues and Anatomy of Root, Shoot and Leaf23-60Unit-3-Structure of Vascular tissues61-84Unit-4-Normal and Anomalous growth85-112BLOCK-2 EMBRYOLOGYPAGE NO.Unit-5- Male Gametophytes114-137Unit-6- Female Gametophytes138-163Unit-7- Fertilization and Post Fertilization164-196BLOCK-3 MORPHOGENESISUnit-8- Plant Morphogenesis and Morphogenetic factorsPAGE NO.198-222Unit-9- Plant Growth Regulators223-243Unit-10-Physiology of Flowering244-270UTTARAKHAND OPEN UNIVERSITYPage 4


ANATOMY, EMBRYOLOGY AND ELEMENTARY MORPHOGENESISBSCBO-202UNIT-1 TOOLS AND TECHNIQUES IN PLANTANATOMY1.1 Objectives1.2 Introduction1.3 Tools in plant anatomy1.4 Techniques in plant anatomy1.5 Summary1.6 Glossary1.7 Self Assessment Question1.8 References1.9 Suggested Readings1.10 Terminal QuestionsUTTARAKHAND OPEN UNIVERSITYPage 6

ANATOMY, EMBRYOLOGY AND ELEMENTARY MORPHOGENESISBSCBO-2021.1 OBJECTIVESAfter reading this unit students will be able to familiar with the history of microscopy and different parts of compound microscopes. to learn different techniques of anatomy like sectioning and staining. to know Mounting media and mounting techniques. to explain the common stains for plant cells1.2 INTRODUCTIONAs in all experimental sciences, research in plant anatomy depends on the laboratorymethods that can be used to study cell structure and function. Many important advances inunderstanding cells have directly followed the development of new methods that have openednovel avenues of investigation. An appreciation of the experimental tools available to the cellbiologist is thus critical to understanding both the current status and future directions of thisrapidly moving area of science. The elements of the plant cell are the membrane and theprotoplast. The protoplast includes the cytoplasm, the nucleus, the plastids, the mitochondria,and other organelles.In the past, the chief objects of study in plant anatomy were the vegetative organs (stem, root,and leaf); today, attention is also given to the structure of flowers, fruits, and seeds. Within thefield of plant anatomy there is:(1) Physiological plant anatomy, which is concerned with the links existing between plantstructure and internal processes.(2) Ecological plant anatomy, which is the study of environmental effects on plant structure.(3) Pathological plant anatomy, which is the study of the effect of disease-producing agents of abiological, physical, and chemical character on plant structure, and(4) Comparative or systematic plant anatomy, which introduces the comparative study ofrepresentatives of the different systematic groups (taxa) - species, genera, families, and soforth for the clarification of their phylogenetic bonds.The basic method used in plant anatomy, or the study of internal plant structure, is thepreparation of thin slices which are studied microscopically. From this the science “derives itsname (in Greek, anatome means “dissection”). The emergence of the field of plant anatomy isclosely related to the invention and perfection of the microscope. The English physicist R. Hookeobserved in 1665 the cellular structure of thin slices of cork, elder pith, and wood from variousplants, using a microscope of his own improved design.The real founders of plant anatomy, however, are considered to be the Italian biologist M.Malpighi and the English botanist N. Grew, who published the first (1675–79) and the secondUTTARAKHAND OPEN UNIVERSITYPage 7

ANATOMY, EMBRYOLOGY AND ELEMENTARY MORPHOGENESISBSCBO-202(1682) works on this subject; in these works the results of a systematic microscopic study ofplant material were presented. Further development came only at the beginning of the 19thcentury. The German scientist J. Moldenhawer in 1812 and the French researcher R. Dutrochetin 1824 were able to divide plant tissue into its component cells through maceration (soaking). In1831 the English botanist R. Brown observed the cell nucleus; this achievement, in combinationwith the studies of the German botanist M. J. Schleiden, played a great role in the founding ofcellular theory, whose author was the German biologist T. Schwann (in 1839). Greatcontributions to the field of plant anatomy were made by the French biologist Edward. vanTieghem and the German biologists Antony de Bary, Carl Von Nageli, K. Sanio, J. Hanstein, andS. Schwendener.1.3 TOOLS IN PLANT ANATOMYThe theoretical knowledge is incomplete without the practical work. Plants are easilyavailable material for the lab studies and their study in the lab adds immense knowledge to thesubject. The practical work develops the scientific outlook and makes the rational approachbased on facts and figures. For a better observation and defining the anatomical features of theplants in the laboratory we use different tools and techniques.Practical Microscopy: The cells of plants are quite minute and microscopic in size, so cannot beobserved by naked eyes. Such objects are visible only under microscopes. Our eye has limitedmagnification or resolution power so unable to distinguish the objects smaller than 0.1 mm.Moreover the living cells are transparent in ordinary light and cannot be distinguished amongvarious cellular components. The microscopes are the most important tools in the plant anatomyand their magnification power is achieved by lenses of various type. The fascinating world ofmicroorganisms and different anatomical features would have remained unknown had themicroscope not been invented.Roger Bacon (1267) described a lens for the first time. However, his observation was notpursued immediately thereafter. In 1590 glass polishers Hans and Zacchrius Jensen constructed acrude type of simple microscope by placing two lenses together, which permitted them to seeminute objects. In 1609-1610 Galileo made the first simple microscope with a focusing deviceand observed the water flea through his microscope. In 1617-1619 the first double lensmicroscope with a single convex objective and ocular appeared the inventor of which wasthought to be the physicist C. Drebbel. This microscope was used to study the cells, plant andanimal tissue, and also the minute living organisms. Till then, the name microscope had not beengiven to this device; the name „microscope‟ was first proposed by Faber in 1625. The credit ofdeveloping a compound microscope with multiple lenses goes to Robert Hooke (1665) ofUTTARAKHAND OPEN UNIVERSITYPage 8

ANATOMY, EMBRYOLOGY AND ELEMENTARY MORPHOGENESISBSCBO-202England. It was only after 1670 that a cloth merchant of Delft (Holland), Antony vanLeeuwenhoek (1632-1723), started his hobby of making microscopes. Considerable progress wasmade in improving the microscope in nineteenth century.Compound Microscope: A compound microscope is the primary tool in the anatomy.Therefore, a clear understanding of structure, use and manipulations of a compound microscopeis a must for all students of anatomy (Fig.1.1).a. Essential parts: The essential parts of usually used monocular compound microscope are thefollowing:Fig. 1.1: Compound MicroscopeLenses: The eye piece with different magnifications (5-20 times). It has field lens towards theobject and eye-lens close to the observer's eye. The objectives generally with three differentmagnifications viz., low (10 X), high (40 X) and oil-immersion (97 X). The focal lengths ofthese are 16 mm, 4mm, and 1.6 mm respectively. These objectives are mounted on a revolvingnose piece for convenience. The eye piece and objectives are fitted at the two ends of a hollowtube called the „body tube‟.Adjustment of objective lens: In some microscopes coarse arid fine focusing adjustment knobsare both provided in order to lower or raise the body tube with lenses for rendering image clear.This is done by rotation of the knobs. The coarse adjustment is meant to bring the object intovision whereas the fine adjustment is used for focusing finer details.UTTARAKHAND OPEN UNIVERSITYPage 9

ANATOMY, EMBRYOLOGY AND ELEMENTARY MORPHOGENESISBSCBO-202Stage: The object to be observed is kept on a glass slides and placed on the stage. It may haveclips to keep the slide in desired position or a mechanical stage for horizontal movement of theobject. In some microscopes the stage may be raised or lowered with coarse and fine adjustmentsfor focusing the object.Mirror: The mirror reflects light, which is transmitted through the object for observing it. Themirror has two planes, one concave and the other plane. When natural light is available the planemirror may be used for reflection of light because concave mirror would form window images.However, with artificial illumination, the concave mirror is necessary for higher magnificationwhereas for lower, the plane mirror may be used.Substage diaphragm: This is meant to control the amount of light transmitted through theobject.Substage condenser: The substage condenser consists of convex lenses which concentrate andintensify the light reflected by-the mirror. With objectives of magnification exceeding 10X, theuse of condenser becomes necessary for narrowing the core of transmitted light, which would fillthe smaller aperture of the objective. The condensers usually employed are called „Abbe‟condensers and these are used with plane mirrors.1.4 TECHNIQUES IN PLANT ANATOMYSolid material should be sectioned in several planes in order to discover the distribution ofthe various tissues within it. The complete investigation of axial structures, such as stem or root,normally requires a transverse (cross) section at one, or more, levels; and radial longitudinal, andtangential longitudinal sections at different depths from the surface to the center. Foliarstructures generally require transverse, and paradermal sections; and vertical longitudinalsections may occasionally be necessary. For anatomical study different techniques are used forvisualizing the cells. Some of the techniques are given below.i) Epidermal peels: The superficial tissues of many plant parts (especially leaves) may bepeeled away in strips thin enough for microscopic examination. To make such a peel, break orcut the surface of the plant apart. Then, grip the epidermis with forceps at one of the cut edges,and pull the outer tissue layer back away from the cut. The resulting epidermal peel should bemounted in water containing a wetting agent, or in alcohol if it is very hydrophobic.ii) Macerations: The three-dimensional form of a cell is most easily seen when the cell isseparated from the surrounding cells of the tissue. Macerating fluids accomplish this througha hydrolysis of the middle lamella. The following method is a gentle, but effective technique:Cut small pieces of the tissue into a mixture of 1 part Hydrogen peroxide, 4 parts distilledwater, and 5 parts glacial acetic acid. Cook the mixture in a 56-60 degree oven for 24 hours.UTTARAKHAND OPEN UNIVERSITYPage 10

ANATOMY, EMBRYOLOGY AND ELEMENTARY MORPHOGENESISBSCBO-202If further macerating is needed, replace the old fluid with a fresh mixture and cook the tissuesfor another 24 hours. Repeat the process until the material is mostly colorless, and may beeasily teased apart with a dissecting probe. When the maceration is complete, rinse thetissues in water in an uncovered container. Stain in 0.25% Safranin in water and mount indilute glycerin.iii) Squashes: Material can be squashed on a slide for cytological examination. This techniqueis most often used for chromosome counts and examination of mitotic structures. Dissectaway non-meristematic tissues, chop the meristem with a scalpel, place a cover slip over thetissue, place paper towels on the cover slip and apply vertical pressure through your thumb.iv) Free-Hand Sectioning: Material should be kept moist while sectioning. Liquid should bekept on the razor blade, so that the sections float as they are cut. In general, it is inadvisableto take particular care over individual sections. Better results are usually obtained by cuttinga large number of slices rapidly, and sorting out the best ones. Sections of uniform thinnessare usually not necessary. Wedge-shaped slices which taper from opaque, overly thickmargins to ultra-thin edges will show useable areas of the proper thickness. When cuttinglongitudinal sections, it is important to use a short piece of material not much longer thanwide. It is impossible to cut satisfactory longitudinal sections of any considerable length bythe freehand technique. Flexible structures, such as leaves, require some support duringsectioning. Many leaves will yield good transverse, and vertical longitudinal sections if rolledor folded so that 10 or more thicknesses are cut at each stroke. If some extra support isnecessary, the material may be inserted into the cut and of a young carrot which has beenpickled in alcohol. The material and the surrounding carrot tissue are then cut at the sametime. This technique should produce results superior to those of the more classical elderberrypith method. To obtain paradermal sections of a leaf, bend it over a finger and cut smallslices off the curved surface. Sections of dry material should first be soaked in alcohol or hotwater to soften it and remove air from the cells.Fig.1.2: Hand sectioningUTTARAKHAND OPEN UNIVERSITYPage 11

ANATOMY, EMBRYOLOGY AND ELEMENTARY MORPHOGENESISBSCBO-202Steps in Sectioning1. Obtain a new double edge razor blade. To minimize the risk of cutting oneself, cover oneedge of the razor blade with masking tape. Rinse the blade with warm tap water to removetraces of grease from the surface of the blade if necessary.2. Hold the plant material firmly. The material should be held against the side of the first fingerof the left hand (or right hand) by means of the thumb (Fig.1.2). The first finger should bekept as straight as possible, while the thumb is kept well below the surface of the material outof the way of the razor edge.3. Flood the razor blade with water this will reduce the friction during cutting as sections canfloat onto the surface of the blade. Take the razor blade in the right hand (or left hand) andplace it on the first finger of the left hand (or right hand), more or less at a right angle to thespecimen.4. Draw the razor across the top of the material in such a way as to give the material a drawingcut (about 450 in the horizontal direction). This results in less friction as the razor bladepasses through the specimen. Cut several sections at a time. Sections will certainly vary inthickness. However, there will be usable ones among the "thick" sections!5. Transfer sections to water, always using a brush, not a forceps or needle.6. Select and transfer the thinnest sections (the more transparent ones) onto a glass slide andstain.Note: For cross sections, special care should be taken during sectioning to see that the materialis not cut obliquely. During sectioning, a number of sections should be cut at the same time andone should not worry about the section thickness at this time. By slightly and progressivelyincreasing the pressure with the razor blade on the first finger, and simultaneously exertingincreasing pressure onto the specimen by the thumb, a number of sections can be cut withoutmoving the material or the thumb. It is best to start cutting with the razor blade right at thesurface of the specimen rather than against the side of the material. Since the root and stemusually have a radial symmetry, it is usually not necessary that a section should be complete, aslong as it includes a portion of the tissues from the center to the outer edge of the specimen.For delicate and hard to hold specimens such as thin leaves and tiny roots, additional supportcan be used to facilitate hand sectioning. The following methods will allow for the sectioningof thin leaves and small, soft specimens such as roots. Tissue pieces can be inserted into asmall piece of pith such as a carrot root. Once the tissue is firmly in place, the hand sectioningtechnique can be applied. Longitudinal sections are also difficult to obtain by hand withoutsupporting material as small stem and root pieces are difficult to hold with one's finger.However, by cutting a v-shaped notch into the pith support, it is possible to hold the tissuefirmly for free hand sections.UTTARAKHAND OPEN UNIVERSITYPage 12

ANATOMY, EMBRYOLOGY AND ELEMENTARY MORPHOGENESISBSCBO-202Treatment of Sections: The student should become familiar with the use of the followingsimple techniques and apply them in the study of laboratory materials: “Wet” water used forpreparing wet mounts of specimens for general observations with a minimum of trapped air.Dilute glycerin used for preparing wet mounts for general observations when rapid drying of themount is undesirable. Chloral hydrate is used to clear whole structures or sections which are nototherwise transparent. Mount the specimen in a few drops of chloral hydrate under a cover glass.Warm the mount over alcohol lamp until it seems more transparent. Do not allow the fluid to boilviolently. Observe the mount directly, or rinse and remount in glycerin. This treatment renderscell walls visible, but removes most cell contents. Starch grains are dissolved, but crystals ofcalcium oxalate do not dissolve. Aniline blue used to stain callose in the phloem, and to stainnuclei and nucleoli.When cut, sections of fresh material should be placed in water, and those of preserved materialsinto alcohol. However, it may be necessary to place sections of some fresh materials into alcoholto get rid of the air within them. In order to avoid unnecessary handling of the sections, alltreatments should be carried out on the slides on which the sections will ultimately be mounted.When the liquid surrounding the section is to be changed,

Department of Botany, Govt. PG College Dwarahat 2. Dr. Sushma Tamta 5, 6 & 7 Assistant Professor, Department of Botany, DSB Campus, Kumaun University, Nainital . ANATOMY, EMBRYOLOGY AND ELEMENTARY MORPHOGENESIS BSCBO-202 UTTARAKHAND OPEN UNIVERSITY Page 2 3. Dr. Nishesh Sharma 8 & 10 .

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