Environmental Plant Physiology Photosynthesis - Aging. Department of Plant and Soil Sciences

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1 Environmental Plant Physiology Photosynthesis - Aging krreddy@ra.msstate.edu Department of Plant and Soil Sciences

2 Photosynthesis and Environment Leaf and Canopy Aging Goals and Learning Objectives: To understand the effects of leaf and canopy aging on photosynthesis process at leaf and canopy level. Photosynthesis and leaf-level aging. Photosynthesis and canopy-level aging.

3 Senescence, Aging and Death and Agriculture Senescence, aging, and death conceived in the past as inevitable, negative processes (wear and tear phenomenon), but now considered as an internal part of differentiation and development. Leaf senescence is one of the most conspicuous processes that has been studied in the context of plant aging and senescence. The terminal phase of leaf development cannot be described simply as a collection of passive and deteriorative processes during which gradual decline in vital systems takes place.

4 Senescence, Aging and Death and Agriculture Extensive physiological and biochemical studies in the last few decades on leaf senescence have suggested that it is highly regulated and active process, which is characterized by differential and sequential changes in almost every sub-cellular compartment. Leaf aging or senescence has implications on agriculture, affecting crop yield and the shelf life of leafy vegetables.

5 Photosynthesis - Aging Leaf Level

6 Leaf and canopy development and aging process 5-day 12-day Leaf Emerging leaf 2-day 8-day Emergence Canopies Squaring Flowering About to abscise Mature crop

7 Photosynthesis - Aging Leaf Level Deep Inside Unfolding leaf Mature leaf Chloroplasts with limited thylakoid development Chloroplasts with welldeveloped thylakoid

8 Photosynthesis - Aging Leaf Level Deep Inside Chloroplasts contain chlorophyll and proteins, and these are the centers of photosynthesis. As leaves grow in size, these centers of photosynthesizing chloroplasts develop very well-defined thylakoids for optimum photosynthesis. Proteins and pigments that function in the photochemical events of photosynthesis are part of the thylakoid.

9 Photoynthesis and Light Response Curves 50 Maximum net photosynthesis Photosynthesis, µmol m -2 s LUE Light Limited Carboxylation Limited 0 LCP where Pn = Respiration PPFD, µmol m -2 s -1 Light saturation point

10 Aging - Photosynthetic Light-response Curves for Cotton Leaves Photosynthesis, µmol CO 2 m -2 s days 27 days 35 days 40 days PPFD, µmol m -2 s -1

11 Photosynthesis, µmol CO2 m -2 s Leaf Aging and Photosynthesis 30/22 C and 360 µl l -1 CO Leaf Age, d

12 Leaf Aging and Photosynthesis Photosynthesis, µmol CO 2 m -2 s PPFD = 1500 µmol photon m -2 s Leaf Age, days

13 Leaf Aging and Light Utilization Efficiency and Conductance LUE, µmol CO 2 µmol -1 photon Light Utilization Efficiency Conductance Leaf Age, days Conductance, mol H 2 O m-2 s-1

14 Leaf Aging - Photosynthestic Parameters Expressed as Fraction of Maximum (= 20 d) 1.5 Fraction of Maximum Light Utilization Efficiency Photosynthesis Conductance Leaf Age, days

15 Leaf Aging and Leaf Nitrogen 5 4 Leaf N, % Leaf Age, days

16 Photosynthesis - Leaf Nitrogen Leaf N, % Photosynthesis, mg CO 2 m -2 s -1

17 Leaf Aging - Leaf Pigments Chlorophyll and Carotenoids Chlorophyll, µg cm Chlorophyll Carotenoids Carotenoid, µg cm Leaf Age, days

18 Leaf Aging - Starch and Sugars Starch 5 4 Sugars, %(w/w) Sugars Starch %(w/w) Leaf Age, days

19 Leaf Aging and Photosynthesis As leaves grow in size, leaf net photosynthesis increases rapidly from leaf unfolding (about zero at leaf unfolding) until the leaf reaches its potential maximum size. Soon thereafter (from about 20 days from unfolding), leaf net photosynthesis declines linearly. Light saturation occurs at lower light levels as leaves age. The onset (from about 20 days) and magnitude (rates) of declines in leaf conductance and light utilization efficiencies are closely coupled with leaf maximum photosynthesis rates. Leaf transpiration rates closely followed photosynthetic rates throughout the leaf development. Leaf pigments (chlorophylls and carotenoids) decline from about 35 days from leaf unfolding, at least in cotton.

20 Leaf Aging and Photosynthesis The concentration of CO2 inside the leaf (Ci), and the number of chloroplasts in the mesophyll remain nearly constant during the aging process, except in the oldest leaves that are about to abscise. Total leaf protein content, RuBP carboxylase and electron transport activity decline parallel to photosynthesis during the aging process. Leaf N also declines from about 20 days, but the decline was not as dramatic as the photosynthetic parameters. The levels of starch and sugars also decrease during the aging process suggesting the loss of photosynthetic activity. Therefore, the physiological deterioration of leaf photosynthetic activity during the aging process may be more related to declining Rubisco protein (photosynthetic capacity) and photosynthetic efficiency (LUE) than to leaf N, leaf pigments and number of chloroplasts. The leaf N and pigments also play a role at a latter stage.

21 Leaf Aging and Photosynthesis Varying light levels during aging by partial shading the leaf did not change leaf photosynthetic characteristics (LUE and Pmax.).

22 Leaf Aging and Photosynthesis The shape and magnitude of photosynthetic capacity and efficiency did not change due to leaf location or position when measured at the same age, at least in cotton. Leaf Aging and Photosynthesis Photosynthesis, µmol CO2 m -2 s /22 C and 360 µl l -1 CO Leaf Age, d

23 Leaf Aging and Photosynthesis In cotton, Pmax is out of sequence with carbon requirements of bolls: 1. Mainstem leaves reach peak rates several days before flowering on the branch and rates are substantially reduced during bollfilling. 2. The branch leaves subtending flower/boll reach maximum photosynthesis (Pmax) about the time of anthesis. 3. Carbon appears to be translocated from remote sites to the developing bolls. Branch leaf Mainstem leaf Flower-bud

24 Photosynthesis - Aging Canopy Level

25 Leaf and canopy development and aging process 5-day 12-day Leaf Emerging leaf 2-day 8-day Emergence Canopies Squaring Flowering About to abscise Mature crop

26 Photosynthesis and Light Response Curves Canopy Photosynthesis, mg CO 2 m -2 s Light Limited Maximum net photosynthesis LUE Carboxylation Limited Light saturation point LCP where Pn = 0 Respiration PPFD, µmol m -2 s -1

27 Canopy Aging - Photosynthesis Photosynthesis, mg CO2 m -2 s ambient Temp. PPFD 1200 µmole m-2 s-1 Flower Open Boll 360 CO Days after Emergence

28 Photosynthesis - Canopy Growth and Aging Canopy Light Utilization Efficiency Canopy Light Utilization Efficiency (µmol CO2 µmol -1 photons) Flowering Boll opening N - Sufficient Days after Emergence

29 Canopy Conductance, mm s -1 Ptotosynthesis - Canopy Growth and Aging Canopy Conductance Flowering Boll opening Days after Emergence N - Sufficient

30 Photosynthesis - Aging Factors controlling aging and photosynthetic process

31 Photosynthesis, mg CO2 m -2 s Canopy Aging - Photosynthesis Response to Carbon Dioxide PPFD 1200 µmole m-2 s-1 Flower Open Boll 1995 ambient Temp. 720 CO2 360 CO Days after Emergence

32 Stem Elongation, cm plant -1 d Boll-load and Vegetative Growth Stem Elongation vs Boll Numbers Linear, y = * X, r ² = Bolls, no. plant -1

33 Canapy Growth - Aging - Daily Net Photosynthesis Seasonal Trends1995 Net Photosynthesis, g CO 2 m -2 d Ambient Temp Days after Emergence

34 Canopy Growth - Aging - Seasonal Trends Response to Nitrogen Nutrition Canopy Photosynthesis, mg CO2 m -2 s N-deficient treatment imposed N-Sufficient N-Deficient Flowering Boll opening Days after Emergence

35 Canopy Aging and Photosynthesis Canopy Light Utilization Efficiency Canopy Light Utilization Efficiency (µmol CO2 µmol -1 photon) Day of N-deficient treatment N - Sufficient Flowering Boll opening N - Deficient Days after Emergence

36 Canopy Aging and Canopy Conductance Response to Nitrogen Nutrition Canopy Conductance, mm s N-deficient treatment imposed N - Sufficient Flowering Boll opening 2 N - Deficient Days after Emergence

37 Photosynthesis, mg CO 2 m -2 s Photosynthesis - Growth - Aging Response to Water Deficits 2 Trt. Start date Water stressed 1 Rewatered Days after Emergence Well-watered

38 Photosynthesis - Aging Canopy level other species - soybean

39 Canopy Aging and Photosynthesis Soybean

40 Photosynthesis - Aging Canopy level other species Indeterminate vs. determinate plant types

41 Canopy Aging - Photosynthesis Species or Plant type Variation Canopy Photosynthesis, mg CO2 m -2 s Elevated Indeterminate Crop: Cotton CO Flowering 1 Determinate Crop: Sorghum Days after Emergence

42 Canopy Aging and Photosynthesis Aging process at the canopy level appears to be a function of leaf-level processes, but modulated by nutrient supply/demand which in turn affects growth and development of various organs including the younger and more efficient leaves produced at the top of the canopy. Sustained photosynthetic efficiency and capacity of canopies require optimum environmental conditions (water, nutrients, including carbon), to continually initiate new leaves. Any management or environmental factor that limits photosynthesis capacity and efficiency can potentially limit crop yield.

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