PLANT SCIENCES II

1. PLANT SCIENCES II BSC-201

2. Plant Domestication– Historical concepts Search for Food Hunter-gatherer society Primary subsistence method involves the direct collection of edible plants and animals from the wild Obtained most of the food (up to 80%) from gathering rather than hunting. Main Source Wild wheat (from German Einkorn, literally "one grain" or "a grain") • Wild einkorn: Triticum monococcum subsp. boeticum : 2n = 14; A genome • Distribution : n. Syria, s. & w. Turkey, n. Iraq, Iran • Harvest: • hand-stripping + bag: average:2.05 kg/h • reconstructed sickle: average: 2.45 kg/h • 46% by weight of actual grain (threshing with wooden mortar and pestle + wind winnowing) • quantity: > needs for one year • Nutritional characteristics: • Acceptable/ good • Poor milling, baking quality • Advantages of einkorn wheat: • abundant & dependable food production • conserves better than meat • Disadvantages: • Fragile rachis • Hulled seed • Changes during domestication: • tougher axis • threshing ratio: wild: 46% grain > cultivated: 73% • more adaptable • more productive • wider leaves

3. Hunter-gatherer societies tend to be relatively mobile: • to provide sufficient resources in order to sustain their population and the variable availability of these resources owing to local climatic and seasonal conditions. • Their population densities tend to be small in number (10-30 individuals), but these may gather together seasonally to temporarily form a larger group (100 or more) when resources are abundant. Domestication of Plants (food security) helped mankind to establish • First farming societies • develop implements • and houses and towns First Farming Societies (12,000-5,000 years ago) Humans began to deliberately grow crops and domesticate a range of plants Population density increased, 60–100 times greater than hunter-gatherer societies, Because cultivated land is capable of sustaining higher population densities than land left uncultivated. The earliest place known to have lived mainly from the cultivation of crops is Jericho (Jordan River in the West Bank of the Palestinian (Fertile Crescent). . By around 8000 BC this community, occupying a naturally well-watered region, is growing domesticated forms of wheat, soon to be followed by barley. Though no longer gatherers, these people are still hunters. Domestication of Plants and Growing them as Crops resulted in Abundant food and consequently Development of Jericho as the first known town, with a population of 2000 or more. A pioneering agricultural community, surrounded by other tribes dependent on gathering food, offers easy pickings which will need vigorous protection. Jerico has protective walls and a tower

4. The Fertile Crescent:The light of civilization first dawned in the Middle East along what is known by historians as the fertile cresent - a cresent-shaped region stretching from just south of modern-day Jerusalem then northward along the Mediterranean coast to present-day Syria and eastward through present-day Iraq then southward along the Tigris and Euphrates rivers to the Persian Gulf. Initially, the Fertile Crescent was only sparsely inhabited but around 8000 BC, it was in this fertile valley that agriculture was first believed to have been developed. Wild wheat and barley grew in abundunce and tribes of nomad hunters and herders began to settle down along the lush banks of the rivers and became the world's first farmers. Agriculture was the spark which lit the flame of civilization. Farming gave rise to social planning on a larger scale as groups of nomadic tribes settled down and joined co-operative forces. Irrigation developed as the need increased to feed and support growing populations. Soon towns were built to afford comfort and protection for these early settlers. Towns like Jericho, Jarmo, Ali Kosh, Catal Huyuk, Beidha and Hassuna were the basis of a new form of human social organization and became the foundation for the first civilization.

5. Introduction to Plant Improvement • What is plant Improvement? • Selection/ development of plants with enhanced performance than the existing genotype/ plants • Early Plant improvement • No scientific methodologies are involved, only on the basis of field performance • Land Races • Varieties of crop plants whose genetic composition is shaped by household agronomic practices and natural selection pressure over generations of cultivation • Systematic Plant Improvement • Improvement in heredity of crops and Production of new crop cultivars which are far better than the existing ones

6. This Systematic Plant Improvement is also referred to as PLANT BREEDING Plant Breeding • is the art and science of improving the heredity of plants for the benefit of the mankind • The goal of Plant breeder is to change the heredity of plant in ways that will improve plant performance

7. Why do we need plant breeding? Performance of CLCV Susceptible Variety S-12 Over The Years at PSC, Farms (Tenant Area) Yield (Mds/Ac) Yield (Mds/Ac) (Direct Area)

8. Plant Breeder Should Know The person involved in Plant improvement should have efficient knowledge of • Needs of the growers and consumers • Characteristics of the crop to be improved, including its wild relatives • Principles of Genetics & Cytogenetics Principles • Special techniques adapted from related fields for the solution of particular problems • Principles of field plot techniques • Principles involved in the design of experiments and the statistical testing of data

9. Related Fields • Botany • Genetics • Plant Physiology • Plant Pathology • Entomology • Agronomy • Plant Biochemistry • Statistics • Computer Sciences • Wheat variety with high yield, disease resistance, Aphid free, less moisture requirement, fertilizer responsive, early maturing, high quality

10. History of Plant Breeding Selection Pre-History • Improvement of wild wheat • Improvement of wild Corn • Improvement in wild potato • Improvement of wild rice Before Mendel • Plant is correct unit for selection: Development of cultivars from progeny of single plant • Screening against wilt in cotton • Reproductive systems in plants Mendel • Laws of inheritance • New branch of science Genetics After Mendel • Hybrid Corn • Dwarf Wheat • Dwarf rice • Strain building (synthetic) in forage Selection/ Introduction Selection/ Introduction Hybridization Tissue culture Biotechnology MAS

11. Basics of Plant Breeding Strategy • Identify the morphological, physiological, pathological traits in a cultivated plant species that contribute to its adaptation, health productivity, and suitability for food, fibre or industrial products • Search out new genes that encode for desired traits in different strains of the cultivated species and their close relatives • Combine genes for the desired traits into an improved cultivar through traditional breeding or new biotechnological procedures • Performance assessment of the improved breeding lines in the local environment in comparison with present cultivars • Distribution of new cultivars that are superior to current cultivars

12. Aims and Objectives of Plant Improvement • Higher yield • Better quality • Shape, size, colour, nutrition, taste, Malting, milling, baking, (cereals) sugar contents in scane, large, fine, strong fibre in cotton, flavour in fruits • Resistance to biotic and abiotic stresses • Crop duration early or late maturity as desired • Growth habit: height, type etc. • Winter hardiness • Lodging resistance • High fertilizer responsiveness • Easier thresh ability • Wider adaptability • Mechanised harvesting

13. Plant Genetic Resources • Plant germplasm • is the genetic source material used by plant breeders to develop new cultivars • Genotypes of particular species, collected from different sources and geographical origins, for use in plant breeding. • Sources of Plant germplasm • Wild relatives • Land races & primitive cultivars • Obsolete cultivars • Advanced breeding lines & other products of plant breeding program • Current cultivars • Thus the germplasm of a crop may be defined as • The sum total of hereditary material i.e. all the alleles of various genes, present in a crop species and its wild relatives

14. Center of Diversity • Area where vast genetic diversity existed for a cultivated crop species (NI Vavilov 1926) Center of Origin NI Vavilov proposed that center of diversity for a crop is center of origin for that crop. • primary centre of origins • Areas where crop plants were domesticated • Secondary Centre of origin • areas where variation continued after domestication

15. Centers of diversity • Chinese Centre • Indian Centre • Indomalayan centre • Central Asiatic Centre • Near Eastern Centre • Mediterranean Centre • Ethiopian Centre • South Mexican and Central American Centre • South American Centre • Chiloe Centre • Brazilian-Paraguayan Centre

16. Agencies engaged in plant breeding • Asian vegetable Research and Development Center (AVRDC) Taiwan • Cabbage, Pepper, tomato, soybean & Mung bean • International center for Agriculture Research in Dry Areas (ICARDA) Syria • Barley, Chick pea, faba bean, tropical forages, lentil & wheat • International center for wheat and maize Improvement (CIMMYT) Mexico • Maize, triticale and wheat • International center for Tropical Agriculture (CIAT) Colombia • Dry beans, cassava, rice and tropical forages

17. Agencies engaged in plant breeding • International Crop Research Institute for the Semi Arid Tropics (ICRISAT) India • Chickpea, millet, peanut, pigeon pea and sorghum • International Institute of Tropical Agriculture (IITA) Nigeria • Cassava, cocoyam, cowpeas, lima bean, maize, pigeon pea, rice, soybean, sweet potato, winged bean and yam • International Potato Center (CIP) Peru • Potato and sweet potato • International Rice Research Institute (IRRI) Phillipines • Rice

18. 3Reproduction in crop plants

19. Flower Structure

20. Kinds of Flowers • Complete: Having all four floral parts i.e. petal, sepal, stamens, pistils e.g. cotton tobacco, potato, cowpeas, soybean, tomato etc. • Incomplete: Lacking one or more floral organs e.g. corn, sorghum, millet, wheat, oat, barley, rice, sugarcane etc. • Perfect: Bisexual, having both stamen and pistil in same flower. Eg. cotton tobacco, potato, cowpeas, soybean, tomato, wheat, oat, barley, rice etc. • Imperfect: Unisexual, having only stamen or pistil in flower • Staminate: having stamen only • Pistillate: having pistil only • Monoecious: having pistillate and staminate on same plant e.g. corn, cassava, castor • Dioecious: Having pistillate and staminate flowers on different plants e.g. hemp, papaya • Cleistogamous: Pollination and fertilization occur before opening of flower, promoting self pollination

21. Nuclear Cell Division • Mitosis (Equational Division) : the daughter nuclei normally receive an exact copy of each chromosome originally present in the nucleus of the parent cell. • is the method of cell division by which new cells are formed in the normal growth and development of plant. • The only form of cell division associated with asexual reproduction • Meiosis Consists of two successive divisions, first reductional (Meiosis I) and second equational (Meiosis II). • Essentially reduces chromosome number from diploid (2n) in the megaspore mother cell to haploid number (n) in gametes. • Associated with sexual reproduction in higher plants

22. Length wise replication of chromosomes to form two identical sister chromatids Appearance of spindle fibres

23. Homologous chromosomes • Chromosomes having genes at corresponding loci controlling a common heredity characteristics Crossing Over • Exchange of chromatid segments Significance of Meiosis • Maintenance of chromosome numbers in plants • Segregation of contrasting alleles leads to their subsequent recombination in following generation • Crossing over provides mechanism for recombination of linked genes and hence new genetic variation.

24. Chromosome number in crop plants Chapter 2 Reproduction in crop plants Table 2.1

25. Types of Reproduction 1. Sexual Reproduction(Union of male & female gametes) • Crops normally Self Pollinated (4 – 5% cross pollination) • Self pollination is the transfer of pollen from an anther to stigma of same flower or to a stigma of another flower on the same plant, or within a clone. • Fertilization resulting from union of male & female gametes produced on the same plant is self –fertilization or autogamy • Crops normally Cross Pollinated • Cross pollination is the transfer of pollen from an anther to stigma of same flower or to a stigma of another flower on a different clone. • Fertilization resulting from union of male & female gametes produced on different clones is cross–fertilization or allogamy • Crops both Self and Cross pollinated • Usually the amount of cross-pollination exceeds that of the normally self pollinated crops, yet does not reach that of the normally cross-pollinated e.g. Cotton ( 5 -25%) pigeon pea (5-40%), Sorghum, tobacco

26. Self Pollinated crops Barley Tomato Bean Triticale Chickpea Vetch Cowpea Wheat Flax Jute Lentil Millet Mungbean Oat Peanut Pea Potato Rice Sesame Soybean Tobacco Cross pollinated Crops Alfalfa Sugar beet Cabbage Sugarcane Carrot Sunflower Castor Sweet potato Cassava Clover Corn (maize) Cucumber Hemp Mustard Onion Pearlmillet Pepper Rye Safflower squash

27. Sporogenesis • Production of microspores and megaspores is known as sporogenesis • Microspores are produced in anthers (Microsporogenesis) • Each anther has 4 pollen sacs with numerous pollen mother cells • Megaspores are produced in ovules (Megasporogenesis) • A single cell in each ovule differentiate into megaspore mother cell Gametogenesis • Production of male and female gametes in microspore and megaspores is known as gametogenesis

28. Male gametogenesis n 2n Second Meiotic Division First Meiotic Division PMC Meiosis n Pollen Microsporogenesis Generative nucleus Nucleus divides n Mitosis in the Generative nucleus n n n Microspore Male Gametes/ sperm Tube nucleus Tube nucleus Microgametogenesis

29. Female gametogenesis n Degenerate n 2n n Second Meiotic Division First Meiotic Division n MMC Meiosis n Megaspore Megasporogenesis Mitosis Mitosis Mitosis Synergid Egg cell n n n v Megaspore Polar Nuclei v v Antipodal Cells Megagametophyte (Embryo Sac) Megagametogenesis

30. Pollen tube sends both male gametes/ sperms into embryo sac • One male gamete/ sperm fuses with egg to form zygote. This process is known as FERTILIZATION. • Second male gamete/ sperm fuses with two polar nuclei. The process is called triple fusion. These three nuclei together form PRIMARY ENDOSPERM NUCLEUS. • The fusion of one male gamete with egg and fusion of second with polar nuclei is called DOUBLE FERTILIZATION. • The fertilized egg develops into EMBRYO • Primary endosperm nuclei divides to form many nuclei covered with cell wall, collectively called ENDOSPERM

32. Asexual reproduction (No union of sexual gametes) Plants may develop either from vegetative parts of plants (Vegetative reproduction) OR Plants may arise from embryos that develop without fertilization (APOMIXIS) Vegetative Reproduction • Propagation through vegetative plant parts (roots, tuber, stolons, rhizomes, stems, leaf cutting or tissue culture) Significance of Vegetative Reproduction • Desirable plant may be used as a variety as there is no danger of segregation • Mutant buds/ branches or seedlings, if desirable can be multiplied and used as varieties. • However it does not allow transfer of genes from one variety to another variety

33. Apomixis (without mixing) In APOMIXIS, seeds are formed but the embryo develops without fertilization. The resulting plants are identical to the parent plant. APOMIXIS may be • Facultative Producing strictly maternally similar plants, • Obligate Reproducing only through apomixis, Obligate apomixis may be • Vivipary: formation of plantlets instead of flowers, no seed is formed • Agamospermy: Formation of seed without the union of egg and sperm nuclei. Agamospermy may be • Adventitious embyony: Embryo develops directly from vegetative cells of ovule and it does not involve production of embryo sac e.g. mango, citrus etc. • Gametophytic Apomixis: Embryo develops without fertilization from egg cell. Gametophytic Apomixis may be • Apospory: Unreduced embryo sac is produced from vegetative cells of the ovule • Diplospory: Unreduced embryo sac is produced from Megaspore Mother Cell Diplospory may be • Parthenogenesis: Embryo develops from the embryo sac without pollination • Pseudogamy: Pollination is necessary for embryo development but fertilization of egg cell does not take place. Fertilization of the secondary nucleus takes place for endosperm development. The endosperm can arise autonomously (Autonomous Apomixis) OR It can arise after fertilization (Pseudogamy)

34. Significance of Apomixis • Apomixis allows plant breeder to fix heterosis • Apomixis allows for rapid multiplication and release of variety Identification of Apomixis • When progeny is identical to mother plant • Through flow cytometery (Flow cytometer is an instrument that can accurately measure the DNA contents of thousands of nuclei

35. Variation in Chromosome Number

36. Chromosomal variation Variation in chromosome may be of two types 1. Variation in chromosome number • 1.1. Euploidy/Polyploidy • 1.2. Aneuploidy • 2. Variation in chromosome structure • 2A. Change in the amount of genetic information • deletions • duplications 2B. Rearrangement of gene locations • inversions • translocations

37. 1. Variation in Chromosome Number Genetic variability forms the basis of plant improvement and variation in chromosome number adds to genetic variability 1.1. EUPLOID: Chromosome number is changed to exact multiple of the basic set Polyploids are euploids in multiple of basic set of chromosome • Diploid 2x • Triploid 3x • Tetraploid 4x • Pentaploid 5x • Hexaploid 6x • Septaploid 7x • Octoploid 8x • EUPLOIDS may be • AUTOPLOIDS: Having Duplicate genome of same species • Autotetraploid: Having Duplicate genome of same diploid species • ALLOPLOIDS: Having Duplicate genome of different species Allotetraploid or amphidiploid: Having Duplicate genome of different species 1.2. ANEUPLOID: Chromosome number of basic chromosome set is changed by addition or deletion of specific chromosomes

38. Ploidy Levels in Different crops

39. Induction of Ploidy • Natural Induction: May arise from • Unreduced gametes: chromosome number is not reduced during meiosis • Natural wide crossing following chromosome doubling • Artificial Induction: • Environmental Shock • Chemical • Colchicine: acts by dissociating the spindle and preventing migration of the daughter chromosomes to poles • It is applied to meristemetic tissue, germinating seed, young seedling, root • Its action is modified or affected by temperature, concentration, and duration of treatment

40. Significance of Polyploidy in Plant Breeding • Permits greater expression of existing genetic diversity • Helps to change the character of a plant by altering number of genomes consequently changing dosage of alleles related to particular trait • Ployploids with uneven number of genomes (Like Triploid and Pentaploids) may result in infertility. This loss of seed production can be used to produce seedless watermelons and banana • About 1/3 to ½ of angiosperms are polyploids • About 70% of wild species of grasses and 23% of legume family are polyploids • Most of the natural polyploids are alloploids • All species don't exhibit vigor with increase in ploidy • Optimum ploidy level for corn is diploid as compared to tetraploid • Optimum ploidy level of banana is triploid (Seedless) • Blackberry is insensitive to ploidy level

41. Artificially Induced Autoploids • Easy to produce: AA doubled to AAAA • Characterized with thicker vegetative parts, increased flower size, less fertile Characteristics of Autoploids • Genetic ratios are simpler in allotetraploid as compared to autotetraploids. Eg. A and a alleles will result in 3 classes (AA, Aa, aa) in alloploid whereas it will result in 5 classes in case of Autoploid • AAAA = quadruplex, AAAa= triplex, AAaa=duplex, Aaaa=simplex, aaaa=nulliplex) • Consequently recessive combinations are less in autoploids forcing breeders to grow larger population Autoploidy often result in • Increased size of meristemetic and guard cells • Decrease in total number of cells, • Reduced growth rate • consequently delayed flowering Use of Autoploids • Due to more vegetative growth autoploids are more suitable in crops harvested for vegetative parts (Forages, root crops, vegetables, flowers) • Useful vigour is obtained by doubling chromosome contents of diploid with low chromosome numbers • Autoploids from cross pollinated crops are more successful than self pollinated crops • Bridging of ploidy levels in interspecific crosses • Diploid treated with colchicine to produce Autotetraploid X tetraploid species

42. Artificially Induced Alloploids • Difficult to produce A, B first need to be hybridized and then doubled to form AABB • After chromosome doubling chromosome from A genome pair with it’s A genome homolog and B with B genome, with no homoeolog pairing between A and B genome. • Homoeolog pairing is restricted by certain genes in natural alloploids like, In wheat, Ph1 present at long arm of 5B chromosome inhibits pairing of homoeolog chromosomes from A and B genomes. • Chromosomes originating from different but similar genomes (like A & B in wheat are different but similar being part of one species) are said to be homoeolog chromosomes (having genes and arrangement of genes in common)

43. Uses of Alloploidy • Identifying genetic origin of polyploid plant • Producing new plant genotypes and plant species • Production of Hexaploid (AABBRR) and Octoploid (AABBDDRR) triticale from rye (RR) and tetraploid (AABB) and Hexploid (AABBDD) wheat • Facilitated transfer of genes from related species • Production of synthetic hybrids of wheat • Fibre strength in cotton • Arboreum(AA) X Thurberi (DD) chromosomes doubled to produce Allotetraploid which was further crossed to hirsutum (AADD) • Facilitating transfer or substitution of individual or pair of chromosomes • IB can tranlocate with 1R chromosome of rye (due to homoeolog)

44. Variation in Chromosomal Number 1.2. Aneuploidy: Chromosome number of basic chromosome set is changed by addition or deletion of specific chromosomes • Commonly results from nondisjunction during meiosis • Monosomy, trisomy, tetrasomy, etc. • Klinefelter and Turner syndromes are examples involving human sex chromosomes • Chromosome deletion lines • Nullisomy - loss of one homologous chromosome pair; 2N – 2 • Monosomy – loss of a single chromosome; 2N – 1 • Chromosome addition lines • Trisomy – single extra chromosome; 2N + 1 • Tetrasomy – extra chromosome pair; 2N + 2

46. Nondisjunction

47. Substitution Line: • Exchange of chromosome between cultivars of same species • Wheat substitution lines Alien Substitution Line: • Exchange of chromosome between cultivars of different species • Wheat X Rye resulting in triticale

48. HAPLOIDY • HAPLOIDS: Plants having gamete number of chromosome • Occur in nature in very low frequency • In many species like corn, wheat, sorghum, barley, rye rice, flax, tobacco, cotton etc. • Can be differentiated from normal diploids (due to smaller size) • Haploidy can be efficiently confirmed by flow cytometery • Haploidy can be less efficiently confirmed by chromosome counting • Haploid plant can be made diploid by treating with colchicine

49. Procedures for Haploid Production • Identification and doubling of naturally occurring haploids • In corn 1/1000 grains is a haploid that arise through development of an unfertilized egg into an embryo by parthenogenesis. Such haploid when doubled is far homozygous inbred line as compared to the one made through successive selfings • Interspecific or intergeneric hybridization followed by elimination of the chromosomes of the wild or distantly related species • In barley H. Vulgare, Rice, wheat through H.bulbosum • In wheat through maize • In wheat through pearl millet and others • Anther or pollen culture • In wheat

50. Use of Haploids Doubling of chromosomes results in diploids that are completely homozygous • This homozygosity achieved in one step is of higher level than normally achieved after 6 generations of selfing • Recessive mutants can be observed at very early stage • Selection of dominant alleles is facilitated • Suitable for mapping populations development