TRANSPIRATION https://www.online-sciences.com/wp-content/uploads/2015/01/plant-12.jpg Science online by Heba Soffar is licensed under a Creative Commons Attribution 4.0 International License. Based on a work at /https://www.online-sciences.com/. TRANSPIRATION • Process where water moves in liquid form in plants, and released in vapor form through aerial parts, but mostly in leaves, to the atmosphere • Energy dependent process • The transformation of liquid to gas phase involves use of energy • 97-99.5% of water taken up is lost in transpiration H2O(liquid) 540 cal g-1 H2O(gas) Importance of Transpiration 1. Keeps cells hydrated 2. Maintains favorable turgor pressure for the transport of nutrients absorbed by the roots from the soil 3. Cools the plant – – heat load is dissipated in the process due to the high heat of vaporization of water If transpiration is extremely high dehydration and desiccation death *** daily water loss – – – large, well-aerated, tropical plant: 500 L corn plant : 3-4 L day-1 (99% of the water absorbed by a corn plant) during its growing season is lost in transpiration) tree-size desert cactus loses less than 25 mL day-1 Types of Transpiration Based on the avenue of exit of water vapor ❑Cuticular transpiration ❑ Loss of water through cuticle ❑5-10% of the water loss ❑Lenticular transpiration ❑Lenticels - pores in the outer layer of a woody plant stem ❑In deciduous species (trees which sheds off leaves) and in some fruits, water loss through lenticels may be quite substantial. ❑Stomatal transpiration ❑Through the stomata ❑As much as 90% of the water lost from plants. What affects diffusion of water from leaf to atmosphere? ❖ Relative humidity (RH) (%) ❖ actual water vapor in the air: water vapor pressure in leaf ❖ In leaves 100% RH; in atmosphere, RH rarely exceeds 90% ❖ water diffuses out from the plants to the atmosphere ❖ Vapor pressure deficit (VPD) (pascal (Pa)) ❖ Actual water vapor pressure - water vapor pressure at saturation at the same temperature ❖ when VPD is 0 Pa (i.e. when RH of the atmosphere is 100%), there is no net movement of water ❖ when the RH of the atmosphere is low, the VPD is high, and the rate of transpiration is faster Soil-Plant-Air Continuum of Water 1. Movement of water from the soil to the root xylem a. Extracellular or apoplastic route - water moves through non-living parts, e.g. capillary spaces of the cell walls and intercellular spaces b. Intracellular route 1) symplastic pathway - plasmodesmata 2) transmembrane or transcellular pathway - vacuolar membrane (tonoplast) and the plasma membranes 2 . Movement of water from root xylem to leaf xylem – transpiration-cohesion-adhesion theory 3. Movement of water from leaf xylem to the air – influenced by RH and VPD – Towards lower water potential (Ψ; expressed in megaPascal, MPa) Movement of water from root xylem to leaf xylem The transpiration cohesionadhesion theory 1. water vapor leaves the air spaces of the plant via the stomates 2. this water is replaced by evaporation of the thin layer of water that clings to the mesophyll cells 3. tension (pulling) on the water in the xylem gently pulls the water toward the direction of water loss 4. the cohesion of water is strong enough to transmit this pulling force all the way down to the roots 5. adhesion of water to the cell wall also aids in resisting gravity FeltyRacketeer6 2019. .https://upload.wikimedia.org/wikipedia/commons/8/85/Transpiration_of_Water_in_Xylem.svg Factors that Affect Transpiration I. Plant Factors 1. 2. 3. 4. 5. Efficiency of evaporative surface Efficiency of water absorption. Other surface/stomatal modifications Phytohormones Canopy structure. 1. 2. Edaphic (soil) factors Atmospheric factors II. Environmental Factors • • • • • Light Relative humidity Temperature Wind velocity Oxygen and carbon dioxide concentrations How Plant and Environmental Factors Affect Transpiration • Leaf number: more leaves, more transpiration • Number, size, position of stomata: more and large, more transpiration, under leaf, less transpiration • Cuticle: waxy cuticle, less evaporation from leaf surface • Light: more gas exchange as stomata are open • Temperature: high temperature, more evaporation, more diffusion • Humidity: high humidity, less transpiration • Wind: more wind, more transpiration • Water availability: less water in soil, less transpiration (e.g. in winter, plants lose leaves) TRANSLOCATION • If TRANSPIRATION is the transport of water and nutrients from soil thru roots and xylem, then TRANSLOCATION is the movement of assimilates (sugars and other chemicals) from the leaf through the phloem to other areas for storage, utilization and consumption by the plant Why need a transport system in plants? • so that cells deep within the plant tissues can receive the nutrients they need for cell processes • In fact: • roots can obtain water, but not sugar, • leaves can produce sugar, but can’t get water from the air CNX OpenStax / CC BY http://cnx.org/contents/GFy_h8cu@10.53:rZu dN6XP@2/Introduction https://commons.wikimedia.org/wiki/File:Figure_ 30_05_07.jpg The vascular tissues: xylem and phloem • Sugars required for metabolism – all the time, in all tissues • Sugars produced only – by source tissues – in light period • Translocation occurs – source to sink over short term – from storage tissues to young tissues over long term Kyla Andrea Mendoza. 2020. Created with BioRender.com Sucrose • is principal photosynthetic product Photosynthetic cell – accounts for most of CO2 absorbed – Glucose, as initial product of photosynthesis, is converted to sucrose which is the major form for transport or translocation • important storage sugar – tap root of carrots and sugar beet (up to 20% dry weight) – and in leaves, eg 25% leaf dry weight in ivy • major form for translocation of carbon – from photosynthetic leaves (source leaves) – in germinating seedlings after starch or lipid breakdown CO2 RuBP Starch PGA Chloroplast 1,3 bisPGA Triose P Sucrose Kyla Andrea Mendoza. 2020. Created with BioRender.com Other compounds are also translocated: Direction of translocation: • From Source: a part of the plant that releases sucrose to the phloem e.g. leaf • To Sink: a part of the plant that removes sucrose from the phloem e.g. root • A plant part can act as source or sink depending on its developmental stage: for example- young leaves act as sink, but later their predominant role would be as source, once they are active in photosynthesis Kyla Andrea Mendoza. 2020. Created with BioRender.com ALLOCATION • The channelling of fixed carbon into various metabolic pathways within an organ or tissue • In a source organ: – Metabolic utilization within the chloroplast – Synthesiszof starch within the chloroplasts – Synthesis of sucrose for export to sink • In a sink organ – Metabolic utilization and growth processes – Storage PARTITIONING • DISTRIBUTION of assimilates to competing sinks – Lower mature leaves feed mainly the roots – Higher mature leaves feed mainly the young leaves and shoot apex • SOURCE LEAVES – Preferentially supply sink organs with which they have vascular connection • Flower or fruit nearest to them (directly above or below them – Basis for flower and fruit thinning Mechanism of translocation of photosynthates 1. Mass or bulk flow (Münch pressure flow hypothesis) 2. Diffusion- slow 3. Cytoplasmic streaming- within the cytoplasm through plasmodesmatal connections between cells 4. Others 1. Facilitated diffusion 2. Active transport across membranes Phloem Cells Kelvinsong / CC BY-SA https://commons.wikimedia.org/wiki/File:Phloem_cells.svg CNX OpenStax / CC BY http://cnx.org/contents/GFy_h8cu@10.53:rZudN6XP@2/Intro duction https://commons.wikimedia.org/wiki/File:Figure_30_05_06.jpg Mechanism of assimilate translocation: The Münch pressure flow hypothesis of assimilate transport from source to sink Alyssa Pham / CC BY-SA https://commons.wikimedia.org/wiki/File:Translocation_ from_the_source_to_the_sink_within_the_phloem.svg APOPLAST PATHWAY • sucrose is loaded into the phloem with the help of active transport. A sucrose transporter protein is used to co-transport H+ and sucrose across the cell membrane. • the apoplast path utilizes ATP to pump H+ against the concentration gradient. • ATP is reduced to ADP+Pi which expels energy that enable H+ to be pushed against the gradient. • The H+ proton that is pumped against the gradient is then used by the sucrose transporter protein to move sucrose through the membrane. • The sucrose accumulated in the companion cell is able to flow down its concentration gradient via the plasmodesmata and into the phloem. Picture cropped and text retyped from work by Emily Xie / CC BY-SA https://commons.wikimedia.org/wiki/File:An_Annotated_Diagram_of_the_Vertical_CrossSections_for_Apoplast_and_Symplast_Pathways_Involved_in_Phloem_Loading.svg SYMPLAST PATHWAY • • sucrose travels in the plasmodesmata which are connections between cells. Therefore, the sucrose is able to flow down the sucrose concentration gradient into the phloem which has a lot of concentration of sucrose. Picture cropped and text retyped from work by Emily Xie / CC BY-SA https://commons.wikimedia.org/wiki/File:An_Annotated_Diagram_of_the_ Vertical_CrossSections_for_Apoplast_and_Symplast_Pathways_Involved_in_Phloem_Loa ding.svg Study questions I. Identify if source or sink 1. Fully expanded mature leaf 2. Unripe fruit 3. Young leaf 4. Flowers II. True or False. If false, give the reason why 1. Transpiration is driven or directed by the amount of water in the soil 2. Water leaves the plants/leaves as water vapor 3. Sugars are the major form of compounds that are translocated 4. When there is RH is 100%, there is 100% transpiration also MINERAL NUTRITION Mineral Nutrition in plants Plants are: Capable of making all necessary organic compounds from inorganic compounds and elements in the environment (autotrophic) Supplied with all the carbon, hydrogen, and oxygen they could ever need (CO2, H2O) Required to obtain all other elements from the soil so in a sense plants act as soil miners. Kyla Andrea Mendoza 2020. Created with BioRender.com Nutrient • any substance that can be metabolized by an organism to give energy and build tissue • growth and development • source of nourishment, especially a nourishing ingredient in a food • providing nourishment Categories of Plant Nutrients • • • Based on Function ❖ Essential ❖ Beneficial Based on amount required by crop ❖ Macroelements ❖ Microelements Based on capability to move from one part of the plant to another ❖ Mobile ❖ Immobile CRITERIA OF ESSENTIALITY 1. If the nutrient is absent, then the plant is unable to complete its life cycle 2. The function of the nutrient must not be replaceable by another element 3. The nutrient must act directly in the metabolism of the plant Functions of the Essential Elements ▪ Structural ▪ important components of biomolecules (e.g. N, P, Ca, Mg, S) ▪ Catalytic ▪ as co-factor of enzymes (e.g. most micronutrients) ▪ Osmotic ▪ regulation of cellular hydration (e.g. K) The Essential Nutrients ▪ Macronutrients: ▪ Nitrogen, phosphorus, potassium, calcium, sulfur, magnesium, oxygen, carbon,hydrogen ▪ Micronutrients: ▪ Iron, boron, copper, zinc, manganese, molybdenum, chloride Macroelements/ Macronutrients Microelements/ Micronutrients Required in relatively large Required in minute quantities quantities like one to 10 milligram like 0.1 mg per gram of dry per gram of dry matter matter Carbon, Hydrogen, Oxygen Phosphorous, Potassium, Calcium and Nitrogen, Sulphur Manganese, Boron, Copper, Molybdenum, Iron, Zinc and and Chloride The essential nutrients- plants take up only INORGANIC nutrients Macronutrient Inorganic Form Micronutrient Inorganic Form Nitrogen (N) NH4+ , NO3- Iron (Fe) Fe+2, Fe+3 Phosphorus (P) H2PO4-, HPO4-2, PO4- Boron (B) BO-3 Potassium (K) K+ Manganese (Mn) Mn+2 Calcium (Ca) Ca+2 Zinc (Zn) Zn+2 Magnesium (Mg) Mg+2 Copper (Cu) Cu+2 Molybdenum (Mo) MoO5- Sulfur (S) SO4-2 Chlorine (Cl) Cl-1 3 Carbon (C) Hydrogen (H) Oxygen (O) Nitrogen (N) Phosphorus (P) Constituent of all organic molecules Component of proteins, enzymes, and nucleic acids In nucleic acids, phytin, coenzymes, adenylases; regulatory function Potassium (K) Osmoregulation; enzyme activator, and protein component Calcium (Ca) Magnesium (Mg) In pectates, and regulatory protein ; regulation of enzyme Integral component of chlorophyll, Mg-ATP; activator of phosphorylation, RuBP carboxylase Sulfur (S) Constituent of several coenzymes, vitamins and amino acids Iron (Fe) Copper (Cu) Zinc (Zn) Components of Fe- and Fe-S proteins, cytochromes, and ferredoxins Activator of several oxidases and lignin synthesis Activator of enzymes Manganese (Mn) Enzyme activator and photosynthetic evolution of oxygen (Hill reaction) Molybdenum (Mo) Enzyme component essential for nitrogenase in bacteria for N2-fixation Boron (B) Complex with protein Chlorine (Cl) Activator of photosystem II; Nickel (Ni) Integral component of urease enzyme Beneficial Elements o Elements which promote plant growth in many plant species but are not absolutely necessary for completion of the plant life cycle o Silicon, sodium, cobalt, and selenium Plant nutrient Source Air Carbon X Oxygen X Water Soil X Hydrogen X Macronutrients Nitrogen X Phosphorus X Potassium X Calcium X Magnesium X Sulfur X Micronutrients Boron X Chlorine X Copper X Iron X Manganese X Molybdenum X Nickel X Zinc X Decline in Soil Fertility ▪ Soil erosion ▪ physical loss and displacement of the fertile topsoil ▪ Geological erosion ▪ Wind erosion ▪ Water-borne erosion ▪ Accelerated erosion due to human activity ▪ Crop removal ▪ Conversion of nutrients to unavailable forms ▪ Formation of insoluble forms ▪ Microbial mediated transformations ▪ Volatilization (especially nitrogen) ▪ Leaching pH affects the growth of plant roots and soil microbes Root growth favors a pH of 5.5 to 6.5 CoolKoon / CC BY https://commons.wikimedia.org/wiki/File:Soil_pH_effect_on_nutrient_availability.svg NUTRIENT DEFICIENCIES • Mineral nutrient deficiencies occur when the concentration of a nutrient decreases below its typical range • Deficiencies of specific nutrients lead to specific visual, often characteristic, symptoms reflective of the role of that nutrient in plant metabolism NUTRIENT DEFICIENCY SYMPTOMS ❑ Chlorosis (uniform or interveinal) or yellowing of the leaves due to chlorophyll degradation ❑ Necrosis (tip, marginal, or interveinal) or death of leaf tissue ❑ Lack of new growth, which may result in death of terminal or axillary buds and leaves, dieback, or resetting ❑ Accumulation of anthocyanin resulting in reddish coloration of leaf tissues ❑ Stunted leaf growth with green, off-green, or yellow color NUTRIENT DEFICIENCY SYMPTOMS P deficient K deficient N deficient N surplus Farnaz Khakpoor / CC BY-SA C deficient https://commons.wikimedia.org/wiki/File:%D9%86%DB%8C%DA%A9% D9%84.jpg ‘Apple Nutrition’ (1941) by Davis, Malcolm B, Hill H, Canada Dept. of Agriculture Retrieved from https://commons.wikimedia.org/wiki/File:Apple_nutrition_(1941)_(19124665773).jpg Patterns of deficiency ▪ Patterns of deficiency are important in identifying which nutrient is deficient or lacking ▪ The location where a deficiency reflects the mobility of a nutrient ▪ Nutrients are redistributed via movement through the phloem ▪ If the deficiency is seen in old leaves = nutrient is mobile ▪ If the deficiency is seen in young leaves =nutrient is fixed or immobile How to address nutrient deficiency? 1. Establish an attainable yield level – the crop’s total needs 2. Calculate the nutrient requirement of the crop 3. Effectively use existing nutrients a. organic fertilizers i. manure ii. plant biomass iii. vermicompost iv. fermented plant juice b. nutrients from irrigation water c. indigenous soil nutrients 4. For deficiencies, top up with or add inorganic fertilizers Plant Growth and Development Development The attainment of size by virtue of growth and architectural style by morphogenesis (differentiation of cells into tissues, organs and organisms) Aspects 1. Growth 2. Differentiation 3. Organization Growth the irreversible increase of cell number, and essentially its dry mass or weight increase in size does not mean an increase in growth. Growth curve • Cells/organs show a definite pattern of growth Differentiation • Reflects the orderly processes by which genetically identical cells become different, forming specialized tissues and organs • The reflection of change in the cell’s biochemical program, controlled by developmental genes Dedifferentiation -the reversal of the cell specialization Important in the repair of injury, where cell near damaged sites become totipotent and reprogram their development Organization • – orientation and integration of differentiated cells in space together with regulated growth whole plant Morphogenesis • the orientation and integration of differentiated cells in space together with regulated growth, and the consequent attainment of form and structure of the complete organism. growth differentiation development • Plants, develop according to a predetermined genetic blueprint • Expression is greatly influenced by signals received from the external environment • Plants are always undergoing development. pattern formation morphogenesis Localization of Growth Essential characteristics of organisms - take up relatively simple substances from environment and synthesize these to complex substances At cellular level - increase in living material leads to increase in cell size and ultimately cell division complex process in multicellular organisms Localization of Growth • Growth is restricted to certain embryonic regions called the meristems Meristems ♣ Where plant growth occurs ♣ Site of repeated cell division of unspecialised cells ♣ Cells differentiate, and become specialised in relation to the function they will perform Basic Structures involved in Plant Growth and Development Embryo Cotyledons, shoots and root apical meristems Meristems - Shoot - Axillary root (apical and lateral) - Floral - Cambiums (cork, vascular) Types of Meristems Apical ♥ tips of roots and shoot ♥ site of primary growth in a plant Lateral ♥ side portions, arising from the cambium (base of nodes and stems), responsible for secondary thickening of the stem and roots Intercalary ♥ inserted between regions of differentiated tissues Types of Growth 1. Indeterminate (ricebean, winged bean) - apical meristems of the roots and stems remain permanently embryonic over long periods 2. Determinate (corn, rice, mungbean) - other plant parts (leaves, flowers, fruits) are embryonic for limited period before the plant reaches maturity, have precise morphology and definite number of parts Internal Growth Mechanisms Correlation Effect The regulatory effects exerted by one part of the plant on the growth or development in another part Internal Growth Mechanisms Organ Differentiation ♥ As shoot increases in bulk, the size of the root system becomes proportionately larger ♥ Reduction in vegetative growth when the plant is fruiting ♥ Stimulation of fruit growth by hormones produced in the developing seeds ♥ Stimulating effects of buds/leaves on the rooting of stem cuttings