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Champter VIII Plant Growth and Movement Key points: Seed germination Plant tissue culture Plant growth Plant movement §1 Seed Germination(种子萌发) Seed germination is the resumption of embryo growth, in general, the radicle goes out the seed coat, depend upon a number of factors. 1. Main factors influencing seed germination: Water, O2, Temp., Light (1) Water soften the seed coat, radicle elongates easily. O2 permeability, embryo metabolism rises. gel to sol activates enzymes. stored macromolecular substance hydrolysis Plant hormone from the conjugate to free form. 30-70% water content for starchy seed germination and more than 110% for protein seed (soybean) germination. (2) Temperature Three cardinal points: “minimum, optimum and maximum” Optimum temperature of germination is the temperature at which seeds germinate fastest with highest germination rate. The larger difference (10℃) between day and night favors to germination. The sowing should be conducted at the temperature 2-3℃ above the minimum temperature in production. (3) O2 Sufficient O2——active metabolism——growth— —fast germination. Deficient O2——anaerobic respiration——over exhausting stored substance and alcohol toxification. Oil seeds (soybean, peanut, sun flower require more O2 than starchy seeds (wheat, corn), RQ<1 。 Shallow-sowing in practice. (4) Light Light seed: germination needs light Dark seed: germinate well under darkness. controlled by red light(660nm) and far-red light (730nm). e.g. tobacco, lettuce, Arabidopsis thaliana e.g. Citrullus (西瓜属) Photophobic seed:germination inhibited by light e.g. eggplant, tomato, onion 2 Biochemical changes during germination Water absorption: rapid→slow→rapid (matrix to osmotic absorption) I: imbibtion; II: physiological changes: III: growth Respiration: From anaerobic respiration to aerobic respiration Organic substances: Starch → glucose Fat → organic fatty acid Protein → amino acids Enzymes: From boundary enzyme → free enzyme De novo synthesis Hormones: From boundary enzyme → free enzyme De novo synthesis 3. Seed longevity and vitality Seed longevity (种子寿命): Time experienced from seed maturation to loss of the germination. most crops 1-3years, lotus seed:1000years, pillow: 12h. Seed vitality (种子生活力)and Seed vigor(种子 活力): Seed viability is the potential ability to germinate Seed vigor is the ability to germinate, grow and produce in the field. It is a complicated term refers to many traits of plants represented by vigor index (Vi). Germination rate (Gr) is percentage of germinated seeds, Gr= number of germinated seeds/ total seedsX100 Vi=SX∑Gt/Dt, where S refers to average fresh weight of shoot or roots; Gt: number of germinated seeds at the day (t); Dt : days of seed germination. Methods to measure seed vitality TTC(氯化三笨四氮唑) BTB(溴麝香草酚蓝) Red ink staining Fluorescence BTB Gelatin Red ink dyeing §2 plant growth and development Development(发育) refers to the sum of all of the changes that an organism goes through in the life cycle-from germination of the seed through growth, maturation, flowering, seed formation, and senescence, it is a qualitative term. Generally speaking, refers to the vegetable to reproduction. Growth(生长) is an irreversible increase in size number and mass. It is a quantitative term. Cell growth consists of division, volume increase and differentiation. Differentiation(分化) refers to qualitative changes that accompany growth. It leads to morphological, structural and functional changes of plants Differentiation occur in various levels Organs Tissues Cells Xylem phloem Molecules Root,stem Conductive Leaf flower fruit Absorptive Mechanic Aerenchyma storage Eye Microscope electron microscope biochemical molecular biology Root hairs… genes,protein Enzyme The Analysis of Plant Growth Plant Growth Can Be Measured in Different Ways Fresh weight Dry weight Cell number Length or height volume Plant Growth Can Be Described in Both Spatial and Material Terms 1. Cell division Factors influencing cell division Hormones: IAA GB CTK Vitamines: VB1, VB2 and Niacin(VB3) stimulate cell division Hormones control the cell cycle in plants 2 Cell elongation Morphology : cell volume increase dramatically as the cell absorbs a great number of water. Combine small vacuoles into a central vacuole so that the nucleus and cytoplasm is pressed to the side toward cell wall. Water is the most important factor for cell elongation, naturally involves a lot of material synthesis and accumulation. •GA, IAA and CTK control cell elongation Cell elongation depends on the elongation of cell wall Components of cell wall Cell wall is composed of : Cellulose Hemi-cellulose Pectin Structural protein Cell wall is a net work 4. Cell Differentiation Keep volume steady and change in morphology and function. Factor affecting differentiation. (1) Plant hormone:IAA/CTK : rooting;shooting (2) Sucrose:lower (1.5-2.5%) sucrose is advantage to xylem differentiation and higher (>4%) to phloem. Middle (2.5-3.5%) to both and the concentration can induce cambium. (3) Light:poor differentiation without light. (4) Polarity : The tissues and organs are arrayed in a precise order along a linear axis that extends from the shoot apical meristem (SAM) at one end to the root apical meristem (RAM) at the other. Upend gives rise to bud and basal end does root. (5) Uneven division of cells. 5. Plant Tissue Culture(植物组织培养) Plant Tissue Culture(植物组织培养), also called micropropagation, is a technique used to propagate cells, tissue, organ and plants under sterile conditions using explants and artificial growth medium, often to produce clones of a plant . The theory basis for plant tissue culture is Totipotency(细胞全能性)refers to all differentiated plant cells retain all the genetic information required for the development of a complete plant. (exception: dead cell or cell without nucleus) Advantages using tissue culture over traditional propagation The production of exact copies of plants that produce particularly good flowers, fruits, or have other desirable traits. To quickly produce mature plants. The production of multiples of plants in the absence of seeds or necessary pollinators to produce seeds. The regeneration of whole plants from plant cells that have been genetically modified. The production of plants from seeds that otherwise have very low chances of germinating and growing, i.e.: orchids(兰花) and nepenthes(猪笼草). To clean particular plant of viral and other infections and to quickly multiply these plants as 'cleaned stock' for horticulture and agriculture Orchids Other Applications Micropropagation can be used to produce large numbers of identical individuals. Micropropagation can also be used to conserve rare or endangered plant species. A plant breeder may use tissue culture to screen cells rather than plants for advantageous characters, e.g herbicide resistance/tolerance. Large-scale growth of plant cells in liquid culture inside bioreactors as a source of secondary products, like recombinant proteins used as biopharmaceuticals. To cross distantly related species by protoplast fusion and regeneration of the novel hybrid. To cross-pollinate distantly related species and then tissue culture the resulting embryo which would otherwise normally die (Embryo Rescue). 1) Culture medium 5 groups of ingredients Minerals such as N P S K Ca Mg Fe Mo Zn Cu B Carbon source: usually sucrose of 2-4% Vitamins: thiamin(VB1), niacin(VB3), inositol,VB6 Organic addictives: such as coconut milk, yeast power. Plant growth regulators: such as 2,4-D, NAA, KT, BA Culture medium can be made as fluid or solid by adding argar of 0.6-0.7%. Culture medium should be sterilized by a autoclave. The optimum temp is about 23-28℃ The optimum light intensity is 500~800mol m-2s-1, generally 600mol m-2s-1 is enough. 2) Explant (外植体) The tissue which is obtained from the plant to culture is called an explant. Theoretically, a totipotent explant can be grown from any part of the plant. However, In many species explants of various organs vary in their rates of growth and regeneration, while some do not grow at all. Growing explants are easily to start cell division. Appendix: ways of getting regenerated plants 1. Callus pathway (organ regeneration pathway) explant→callus→new plant 1) Adventitious roots → → →adventitious bud 2) Adventitious bud → → →adventitious root 3) Adventitious root and bud simultaneously -----------------------Dedifferentiation (脱分化) is a process of regression of a specialized cell or tissue to a simpler, more embryonic, unspecialized form. Redifferentiation (再分化)is a process by which a group of once dedifferentiated cells differentiate to specialized forms. 2. Embryoid pathway: Develop from a somatic embryo (体细胞胚)to a whole plant, just like from zygotic embryo to whole plant. It also experiences “Early globular stage”, “heart stage”, “ Torpedo stage” and “Mature embryo”. Cluster seedling pathway: Use buds as explant §3 Plant growth 1. Grand period of plant growth(植物生 长大周期) Grand period of plant growth refers to that the growth of tissues, organs and even a whole plant exhibit a typical model with “slower-faster –slower” in life cycle. The total growth appears S-shape growth curve---logistic curve. Grand growth period Growth curve Time 2. Factors influencing plant growth 1) Light 2) Temp. 3) Water 4) Minerals 5) others 1) Light Indirect:by photosynthesis and transpiration. (High energy) Direct:photomorphogenesis. (Low energy) High irradiation inhibit plant gowth Blue light, UV inhibit plant growth 2) Temperature Three cardinal points: “minimum, optimum and maximum temperature for growth” Optimum temperature for growth is the temperature at which plant grows fastest but not strongest. Suboptimum temperature for growth is the temperature at which plant grows slower but strongest than at optimum temperature . Difference in temperature between day and night (thermoperiodicity of growth: 生长的温周期 or daily periodicity of growth: 生长的昼夜周期 ) controls plant growth. 3).Seasonal periodicity of growth mainly controlled by temperature, light and water. General speaking, slowly in the Spring, fast in the summer and again slowly in the autumn, stop in the winter. 4).Water, O2 and minerals “Dry for root, wet for bud”. Water, Oxygen, Minerals 3. Plant growth correlation (植物生长的相关性) The growth among the different organs in a whole plant coordinates and inhibits each other, which is called the plant growth correlation. The correlation mainly includes root and shoot, stem and branch vegetable organs and reproductive organs. 1) Root vs. Shoot Shoot provides photoassimilate, IAA and VB1 to roots; Root provides water, mineral nutrition, plant hormones (CTK、GA, ABA) and partial AA to shoot. They must coordinate each other. Root-shoot ratio(根冠比) Root-shoot ratio is an index for the coordinative between root and shoot, expressed as root weight/shoot weight: Factors influencing root-shoot ratio and its regulation (1) H2O soil water content↓,the ratio↑. (2) Regulation with nutrition • N↑, the ratio↓;N↓, the ratio ↑. • Leaf vegetable applied with heavy N. • P and K↑ the ratio ↑. • Tuber root applied with much P and K. (3) Sufficient light ↑ the ratio ↑. (4) Temperature ↓, the ratio ↑. (5) Tilling and breaking roots in the surface of soil,the ratio↑. Main stem and Branch —— Apical dominance(顶端优势) The phenomenon that main stem apices grows fast and axillary bud or branch grows slowly of a plant is called apical dominance . The reasons for dominance: (1)Nutrition hypothesis (K. Goebel,1900); (2)IAA hypothesis---direct-inhibition model (Thamann and Skoog, 1934) , IAA inhibits branch; (3) Nutrition supply and plant hormone (IAA, GA and CTK) control theory Apical dominance of a tree Vegetative Vs. Reproductive growth The better vegetation promotes reproduction and the poorer vegetation results in poorer reproduction . Over-vegetation inhibits reproduction. Overgrowth often gives rise to late flowering, bad setting, fruit falling and yield decreasing. Over-reproduction reduces the vegetative organ development and leads to plant premature, for fruit tree, leads to “big and small year”. Locomotion Movement Animal such as bee In dark Under sunlight §4 Plant movement Tropical movement (向性运动) Nastic movement(感性运动) Rhythm movement(昼夜节律) 1. Tropic movement Tropical movement is the movement caused by unilateral stimulus, e.g. phototropism, gravitropism, chemotropism and hydrotropism. 1) Phototropism Positive phototropism (正向光性): Sunflower Negative phototropism (负向光性): Air roots Diaphototropsm (横向光性):: leaves 2) Gravitropism Positive gravitropism (正向重力性): Roots Orthogravitropism (负向重力性): Shoot Diagravitropism (横向重力性): subterraneous stem Statolith (平衡石)is the amyloplast in the root cap, each amyloplast is composed of 1-8 starch grains. One cell contains about 4-12 amyloplasts. Figure 18.11 Root cap perceive gravity. Removing root cap will not affect growth response but the gravitropic response is abolished. Regeneration of root cap requires gravity and plants can respond to gravity after root cap is regenerated. The columella cells (中柱鞘细胞)are the most sensitive cells for sensing gravity. Figure 18.16 2. Nastic movement Movement is caused by stimulus of no direction. Include: growth movement( irreversible) and turgor movement (reversible) 1) Nyctinasty movement(感夜运动) Maranta (竹芋) at afternoon (left) and in the night (right) 2) Seismonasty(感震运动) The sensitive plant (Mimosa pudica) have special structures (pulvinus) at the bases of petioles and leaflets that allow the leaf parts to close up when touched (arrow). Pulvinus(叶褥) Structure of ventral (腹侧) and dorsal (背侧)motor cells in the pulvinus 3. Circadian rhythm (近似昼夜节奏) or physiological clock (生理钟)or biological clock(生物钟) Some movement is controlled by inherent factors such as such as oxygen evolution and respiration, alternately through highactivity and low-activity phases with a regular periodicity of about 24 hours, this movement is called Circadian rhythm or physiological clock or biological clock. 菜豆叶在恒定条件(微弱光,20℃)下的运动 Figure 19.15 Basic model of circadian clock Figure 19.16 Clock gene (CCA1?)