Mineral_Nutrition-Liao_Hong-1(minimizer)

1、Principles of Mineral Nutrition in Higher PlantsDr. Hong LIAO Department of Plant NutritionSouth China Agricultural UniversityTel: 85283380, Email: Text Book (教课书教课书): 高级植物营养学高级植物营养学 ( (科学出版社，科学出版社，2003)2003)Main Reference Books (主要参考书）主要参考书）: 植物营养与肥料植物营养与肥料 (浙农大，浙农大，1991)农业化学农业化学 (总论总论) (第二版第二版) (中

2、国农大，中国农大，1994)根系生物学：原理与应用（科学出版社，根系生物学：原理与应用（科学出版社，2007）Principles of Plant Nutrition (Third Edition) (K. Mengel, E. A. Kirkby. 1982)Mineral Nutrition of Higher Plants (Second Edition) (H. Marschner. 1995)1. Mineral Nutrition of Plants: Principles and Perspectives (Second Edition) (E. Epstein, A. J.

3、Bloom. 2005)Table of ContentsIntroductionNutrient ElementsFunctions of Some Essential NutrientsPlant Nutrition Under StressesPlant Nutritional GeneticsPerspectivesTable of ContentsIntroductionNutrient ElementsFunctions of Some Essential NutrientsPlant Nutrition Under StressesPlant Nutritional Geneti

4、csPerspectivesWhy Do We Study Plant Nutrition?Population growthFood productionNutrient requirementSource: World Population Prospects: The 2008 Revision, /unpp1950-2050年世界总人口变化及预测(From Chen XP)9.1510.57.9611.0根据联合国预测，按照中等生育率，根据联合国预测，按照中等生育率，20502050年世界人口将达到年世界人口将达到91.591.5亿。亿。China: C

5、hallenges for Production (million tons)Annual growth (percent per yr)1961198020002007 1961-19801981-2000 2001-2007World8771550206023493.8 1.6 1.8 China1102804074587.8 2.2 1.5 India871402352603.0 3.2 1.4 UK101924195.0 1.1 - 2.5USA1642703434153.2 1.3 2.6 Africa46731121372.8 2.6 2.9 Source: FAO，2010, h

6、ttp:/1961-20071961-2007年部分国家及地区谷物产量及年增长速年部分国家及地区谷物产量及年增长速率率新世纪以来，我国粮食年均增长速率低于世界平均水平新世纪以来，我国粮食年均增长速率低于世界平均水平 (From Chen XP) 。Crop Production in China (1949-1994)Manure Manure Manure Manure N N N N Variety P P Pesticide K Variety Pesticide Variety0100020003000400050001940195019601970198

7、019902000(Year)Yield(kg/hm2)Fertilized to death, Nosengo, 2003, NatureOver- Fertilization Causes Environmental PollutionTN：10-29mg/LTP：0.5-2.8mg/LSS：0.6-1.4g/LThis map identifies 415 eutrophic and hypoxic coastal systems worldwide. Of these, 169 are documented hypoxic areas. There are 24 eutrophic c

8、oastal areas with documented hypoxia within Asia. (World Resources Institute, 2007） /map/world-hypoxic-and-eutrophic-coastal-areasImprovements in the water quality of many freshwater and most coastal marine ecosystems requires reductions in both nitrogen and phosphorus inputs.Plant Nutrit

9、ionPrinciples of Plant NutritionFertilizersCrop FertilizationMethodology for Plant Nutrition ResearchAbsorption, translocation, assimilation and mechanisms of nutrient elements (essential or nonessential) of plantsBroad Sense（广义定义）（广义定义）Narrow Sense（狭义定义）（狭义定义）Liebigs Three Theoriesv Theory of miner

10、al elements 矿质营养学说矿质营养学说v Theory of nutrient return 养分归还学说养分归还学说v Law of minimum nutrient 最小养分律最小养分律Justus von Liebig1803-1873Justus-Liebig-UniversityMineral elements are the only sources of nutrients for plant growth in soils. The nutritional roles of manure and other organic fertilizers in plant g

11、rowth are not from their organic matter, but from their decomposed mineral elements. Theory of Mineral Elements （矿质营养学说）（矿质营养学说）Liebigs Three TheoriesBasis of modern soilless culture2. Theory of Nutrient Return（养分归还学说）（养分归还学说）All the nutrients taken by plants have to be retuned to soils. Otherwise,

12、the soil fertility will gradually get down.Basis of environmental and sustainable agriculture Plant growth is limited by the most limiting nutrient as illustrated by barrel staves of varying lengths representing growth-controlling factors2. Law of the Minimum（最小养分律）（最小养分律）Basis of modern fertilizati

13、on Table of ContentsIntroductionNutrient ElementsFunctions of Some Essential NutrientsPlant Nutrition Under StressesPlant Nutritional GeneticsPerspectivesNutrient Elements of Plants植物的营养元素MnBFeSNCOHCaKPCuClZnMgMo Ni Ni Compounds of Plants植物体的组成成分Fresh plant (新鲜植株新鲜植株)75%95% water (水分水分)5%25% dry mat

14、ter (干物质干物质)dried (烘干烘干) 75C 95% evaporated as gas（气体挥发）（气体挥发） 5% ash (灰分灰分)burned (煅烧煅烧) 525CThree Criteria for Essential Mineral Elements必需营养元素的三大标准必需营养元素的三大标准 A given plant must be unable to complete its life cycle in the absence of the mineral element (Essential，必需性必需性)The function of the elemen

15、t must not be replaceable by another mineral element (Specificity，专一性专一性)The element must be directly involved in plant metabolism or must be required for a distinct metabolic step such as an enzyme reaction (Direct，直接性直接性). C、H、O 天然营养元素天然营养元素 非矿质元素非矿质元素Macronutrients 来自空气和水来自空气和水大量元素大量元素 N、P、K 植物营养

16、三要素植物营养三要素(0.1%以上以上) 或或肥料三要素肥料三要素 Ca、Mg、S 也称也称中量元素中量元素 矿质元素矿质元素 来自土壤来自土壤Micronutrients Fe、Mn、Zn、Cu、微量元素微量元素(0.1%以下以下) B、Mo、Cl、NiEssential Mineral Elements 必需营养元素必需营养元素Beneficial Elements of Plants植物的有益元素 某些元素适量存在时能促进植物的生长发育；某些元素适量存在时能促进植物的生长发育；或者是某些特定的植物、在某些特定条件下或者是某些特定的植物、在某些特定条件下所必需的，这些类型的元素称为所必需的

17、，这些类型的元素称为“有益元有益元素素”，也称，也称“农学必需元素农学必需元素”。Sodium (Na), Silicon (Si), Cobalt (Co).CO2O2H2OO2K+Cl-Zn2+Mn2+Mg2+NO3-NH4+PO4-Ni2+Cu2+SO42-Fe3+Ca2+Fe2+BOHOHOHMovementuptakeBasic Processes of Plant NutritionTranslocationNUTRIENT UPTAKE BY THE ROOTIon Transport 离子运输Short distance transport 短距离运输短距离运输Long dis

18、tance transport 长距离运输长距离运输Rhizodermis 根表皮细胞根表皮细胞Cortex皮层细胞皮层细胞Stele中柱中柱Long distance transport 长距离运输长距离运输Xylem Transport：One direction: from root to shoot（从根到叶）（从根到叶）Phloem Transport:Two direction:From source to sink (从源到库）从源到库）Ion Mobility in the PhloemHigh mobility Intermediate mobility Low mobili

19、ty Potassium (K)Magnesium (Mg)Sulfur (S)Nitrogen (amino-N)Chlorine (Cl)Sodium (Na)Iron (Fe)Zinc (Zn)Copper (Cu)Boron (B)Molybdenum(Mo)Calcium (Ca)Manganese (Mn)Table of ContentsIntroductionNutrient ElementsFunctions of Some Essential NutrientsPlant Nutrition Under StressesPlant Nutritional GeneticsP

20、erspectivesFunctions of Some Essential NutrientsNitrogen （氮）（氮）Phosphorus（磷）（磷）Potassium（钾）（钾）Iron（铁）（铁）Functions of Some Essential NutrientsNitrogenPhosphorusPotassiumIronMain N-containing Compounds in Plants植物体内的主要含氮组分植物体内的主要含氮组分1) Proteins and amino acids 蛋白质和氨基酸蛋白质和氨基酸2) Nucleic acids 核酸核酸3) Enz

21、ymes 酶类酶类4) Chlorophyll 叶绿素叶绿素5) Others : Alkaloids, vitamins and phytohormones 其它：生物碱、维生素和植物激素等其它：生物碱、维生素和植物激素等Main Uptake Forms of Nitrogen 氮素的主要吸收形态氮素的主要吸收形态 1) NH4+- N: ammonium-N 铵态铵态-氮氮2) NO3- N: nitrate-N 硝态硝态-氮氮3) Soluble N-containing organic materials可溶的有机态氮可溶的有机态氮: urea, amino acids and am

22、ides4) NH3 and NO2: ammonia and nitro-dioxide(by shoots)5) N2: by biological N fixation 生物固氮生物固氮There is considerable variation in the details of nitrogen assimilation in different plants. Plant roots can import nitrate, a m m o n i u m , a n d o t h e r nitrogenous compounds from the soil. For use

23、in synthesis of amines and amides, nitrate must be reduced to nitrite and then to ammonium. Nitrate reduction in the cytosol and storage in the vacuoles are processes that can occur in either the roots or the leaves. Nodulated plants are able to take up fixed nitrogen from the soil (not shown) but,

24、through the action of symbiotic bacteria, c a n g e n e r a t e ammonium also by reducing N2. The ammonium from nitrogen fixation is assimilated into amino acids and ultimately incorporated into amide amino acid (glutamine or asparagine) or ureides.Ammonium uptake and transport铵态-氮的吸收与运输In general:

25、active transport process, including multiple transport systems. The Km values for the transporters range from 10 to 70 uM for NH4+, including two high-affinity systemsAMT family: plant genes encoding for ammonium transport was identified in tomato and Arabidopsis. NH4+ transport depends on electroch

26、emical potential rather than direct ATP synthesis. AMT type genes have shown wide expression in some cases and root hair-specific expression in others.SAT1: a specific ammonium transporter, localized on the symbiosome membrane in soybean, specifically expressed in nodulated roots. Its Km is about 5

27、mM. Ammonium assimilation ATP ADPPi COOH H2NCH CH2 CH2 C=O OHH (GS) NH2 铁氧还蛋白 2e- 2H+ （GOGAT） COOH CO CH2 CH2 COOH -酮戊二酸 COOH H2NCH CH2 CH2 COOH 谷氨酸 COOH H2NCH CH2 CH2 CO NH2 谷氨酰胺 谷氨酸Nitrate uptake and transportIn general: Nitrate uptake is carrier-mediated, active transport. As nitrate diffuses fro

28、m the soil solution into apoplasm of the root, it is taken up by the epidermal and cortical cells. Plants utilize both high-affinity and low-affinity transport mechanisms. Gene products associated with both high- and low-affinity nitrate uptake have been identifiedIn general: Two nitrate transporter

29、 gene families, NRT1 and NRT2, have been discovered. The NRT1 family: includes nitrate transporters with dual affinity (both low and high Km) or low affinity. The NRT2 family: encodes transporters that contribute to the inducible high-affinity uptake system. Nitrate assimilation by plant cells invol

30、ves transport of nitrate across the plasma membrane and then reduction to ammonia in a two-step process. A proton-pumping ATPase maintains the electrochemical gradient that drives cellular uptake of nitrate. The values shown for electrical potentials and intercellular nitrate concentrations are typi

31、cal but can vary significantly.Overview nitrate assimilationRelative amounts of nitrate and other nitrogen compounds in the xylem exudates of various plant species. The plants were grown with their roots exposed to nitrate solutions, and xylem sap was collected by severing of the stem. Note the pres

32、ence of ureides, specialized nitrogen compounds, in bean and pea (After Pate 1983) .Biological Nitrogen Fixation (BNF)Biological nitrogen fixation accounts for most of the fixation of atmospheric N2 into ammonium, thus representing the key entry point of molecular nitrogen into the biogeochemical cy

33、cle of nitrogen.Free-living and symbiotic bacteria fix nitrogenSymbiotic nitrogen fixationPhoto of root nodules on pea (left) and soybean (right)Symbiosis: Nodules between Rhizobia and Plants Plants: organic acids- supply CBacteria: produce NH3Plant roots (GS-GOGAT system)N2 + 16 ATP + 8e- + 8H+2NH3

34、 + H2 +16ADP +16 PiNitrogenaseDinitrogenase reductase(Fe-protein)Dinitrogenase(MoFe-protein)The General Reaction Catalyzed by NitrogenaseNitrogen DeficiencySmall plants, yellow leaves starting from old leavesRice plants of N-deficiency (left), Yellow colors in old leaves.Peanuts of ?-deficiencyNitro

35、gen recycling on citrate leaves with inadequate N (left)(A) Green terminal leaves (B) Yellowing(C) DefoliationFunctions of Some Essential NutrientsNitrogenPhosphorusPotassiumIronFunctions of Phosphorus in PlantsStructural element: Nucleic acid , phosphlipidsEnergy transfer: AMP, ADP, ATP, UTP, GTP a

36、nd PPiModulate enzyme activitiesRegulatory role of inorganic phosphateSoils Prone to Be P Deficient in the World PiP iR-COO-Fe, Al-PR-COO-PtaseOrganic PPtaseH+Ca-PH+TransporterTransporterArum-typesParis-typesSandy DicksonGallaud 1904, 1905: Smith & Smith 1997Symbiosis: Mycorrhizal Fungi and Plants P

37、hosphorus DeficiencyStunted, reddish purple tips and leaf marginsCornFunctions of Some Essential NutrientsNitrogenPhosphorusPotassiumIronrCompartmentation and cellular concentrationsrEnzyme activationrProtein synthesisrPhotosynthesisrOsmoregulationrCell extensionrStomatal movementrLight-driven and s

38、eismonastic movementrPhloem transportrCation-anion balanceFunctions of PotassiumHigh-affinity K+-transporterLow-affinity K+-transporterBoth high and low -affinity K+-transporterPotassium UptakeK+-transporterHAKHigh affinityK+-H+ co-transporterKUPHigh, High-low-affinityK+-H+ co-transporterHKTHigh aff

39、inityK+-Na+ co-transporterLow-affinity for Na+Na+-Na+ co-transporterLCTLow-affinity for Na+/K+/Ca2+UniporterGeneAffinityAccompany ion Potassium DeficiencyYellowing and necrosis of leaf margins, or brown leaves (fired leaves) starting from old leaves Functions of Some Essential NutrientsNitrogenPhosp

40、horusPotassiumIronFunctions of Iron1. Iron-containing constituents of redox system: Heme, iron-sulfur proteins2. Other iron-containing proteins3. Effects on chlorophyll development and photosynthesis4. Effects on root growthModel of Fe uptake by dicots and non-graminaceous monocots, depicting the ke

41、y plasma membrane-localized components that are induced in roots by Fe deficiency. These include an inducible ferric reductase that reduces Fe(III) chelates in the rhizosphere, releasing Fe2+ for transport across the plasma membrane by a high-affinity Fe2+ transporter. Also shown is the plasma membr

42、ane H+-ATPase, which is induced by Fe deficiency and acidifies the rhizosphere, increasing Fe solubility.Strategy IModel of Fe uptake by graminaceous plants. Phytosiderophores (PS) are synthesized in the cytoplasm from methionine by way of nicotianamine (1). An Fe deficiency-inducible plasma membran

43、e (PM) transporter is involved in release of PS into the rhizosphere (2). Another Fe deficiency-inducible transporter in the PM recognizes the Fe-PS complex and transports it intact into the cytoplasm (3). Phytosiderophoreseffective ferric chelatorsmugineic avenic acidsStrategy IIFe-deficient potato

44、 plant Fe-deficient corn leavesFirst in the younger leaves;Youngest leaves may often be completely white and totally devoid of chlorophyll ;For cereals: alternate yellow and green strips along the length of the leafIron Deficiency(Chlorosis)Table of ContentsIntroductionFunctions of Some Essential Nu

45、trientsPlant Nutrition Under StressesPerspectivesv Biotic stressesv Abiotic stressesPlant Nutrition Under Stresses Acid Mineral Soils Alkaline Soils Waterlogged and Flooded Soils Heat Stress Freezing Stress Saline Soils Nutrient toxicity and deficiencydiseases, insects, weeds etc. Distribution of Ac

46、id SoilsACID SOILS AND ALUMINUM TOXICITYThird most abundant element in the earths crust30 - 40 % Worlds arable soils are acidicAt soil pH values below 5, Al3+ solubilizes into the soil solutionAl Toxicity: Al3+ is the major toxic speciesAl toxicity is an important limit to agricultural productivity

47、worldwideMany cellular targets Cell division and elongation Generates free radicals Al3+ is an analog of Mg 2+Yield reduction inhibition of root growth inhibition of nutrient and water acquisitionAl Toxicity: Al3+ is the major toxic speciesAluminum ToxicityEffects of Al exposure on root growth of Al

48、-resistant Atlas66 and Al-sensitive Scout 66 cultivars of wheat. Seedlings were grown for four days in 0.6 mM CaSO4 solutions containing 0, 5, 20, or 50 uM AlCl3 (pH 4.5). Exclude aluminum release Al-binding ligands Sequester aluminum Store in vacuole Stress responsesB)Commonly secreted organic acids that detoxify aluminum (Al). (a) Root elongation during 20h exposure. (b) Al accumulation (pink color) stained by 0.1% solution of Eriochome cyanine R. The absence of color indicates that the organic acids has chelated the Al and prevented its accumulation in the root apices. The different abili