Avocado Tree Physiology Understanding the Basis of Productivity
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1 Avocado Tree Physiology Understanding the Basis of Productivity R. L. Heath, M. L. Arpaia UC, Riverside M. V. Mickelbart Purdue University Raw Materials Labor Product Light Carbon Dioxide Temperature Water Nutrients Labor Fruit 1
2 Research priorities addressed Canopy management, tree density, tree architecture Development and refinement of a model to predict phenological events for the avocado Innovative practices to increase efficiencies of grove operations and orchard profits Problem: Lowered Productivity in California Photosynthesis is the Production Line of the Factory What factors limit photosynthesis? 2
3 Problem: Lowered Productivity in California Photosynthesis is the Production Line of the Factory What factors limit photosynthesis? STOMATA CO 2 & WATER FLOW lower leaf surface of Hass avocado leaf [CO 2 ] out stomata Light Assimilation [CO 2 ] in Bulk Air Boundary Air Internal Air [H 2 O ] in carbohydrate [H 2 O ] out epidermis leaf 3
4 Problem: Lowered Productivity in California WHY? Several reasons BUT [1] Stomata close-down in error, lowering assimilation Stomata conductance altered by water loss [2] Light is limiting for assimilation, relative to size of canopy Potential solutions: [1] Increase productivity through manipulation of air relative-humidity. [2] Alter the canopy of the trees to change air flow & light absorption Denser canopy to control water loss. Research Plan: [1] Determine the assimilation efficiency upon stomata conductance [2] Determine the variation of conductance upon relative humidity. [3] Build a simplified model of assimilation photosynthetic assimilation (μ mol / m sec) LiCor 62 porometer Field conditions One Limit to Productivity: Stomata stomata control of assimilation stomatal conductance (mmol / m 2 sec) [1] During normal conditions assimilation governed by stomata conductance [2] During most stress conditions stomata conductance declines but assimilation still governed by stomata conductance From Mickelbart ( ) & Xuan ( ) 4
5 Light Intensity Light Intensity Accomplishments to date Light Flecks Leaf area Leaf processes Light intensity and assimilation Model of productivity Sap flow Light Flecks inside canopy Field 1 (measured by light meter on east side, ½ way into canopy, 6 ft off ground) 96 lum/ft 2 64 lum/ft 2 Tree 9-1 6/22 Tree 8-1 6/21 7 min 2 sec 15 lum/ft 2 9 AM Noon 2 lum/ft 2 9 AM Noon 5
6 Speed of stomata change Licor Experiment (Hass Leaf) Declining Intensity Increasing Intensity Conductance (mol/m2 sec) Conductance Intensity Light Intensity (umol/m2 sec) Time of experiment (min).3 2 Licor Experiment (Hass Leaf).25 Declining Intensity Increasing Intensity Decline Conductance (mol/m2 sec) Conductance Intensity Light Intensity (umol/m2 sec) Lower intensity = closed stomata Lower intensity & closed stomata = lower assimilation Lower assimilation & closed stomata = higher internal CO Time of experiment (min) 6
7 .3 2 Licor Experiment (Hass Leaf).25 Declining Intensity Increasing Intensity Decline Conductance (mol/m2 sec) Conductance Intensity Light Intensity (umol/m2 sec) Lower intensity = closed stomata Lower intensity & closed stomata = lower assimilation Lower assimilation & closed stomata = higher internal CO Time of experiment (min) 45 6 ambient CO 2 4 Assimilation (umol/m2 sec) 4 2 Assimilation Internal CO internal CO2 (ppm) respiration Time of experiment (min).3 2 Licor Experiment (Hass Leaf).25 Declining Intensity Increasing Intensity Decline Conductance (mol/m2 sec) Conductance Intensity Light Intensity (umol/m2 sec) Lower intensity = closed stomata Lower intensity & closed stomata = lower assimilation Lower assimilation & closed stomata = higher internal CO Time of experiment (min) 45 6 ambient CO 2 4 Increase Assimilation (umol/m2 sec) 4 2 Assimilation respiration Internal CO internal CO2 (ppm) Higher intensity = higher assimilation Higher assimilation & closed stomata = lower internal CO 2 Lower internal CO 2 = opening of stomata Time of experiment (min) 7
8 .3 2 Licor Experiment (Hass Leaf).25 Declining Intensity Increasing Intensity Decline Conductance (mol/m2 sec) Conductance Intensity Light Intensity (umol/m2 sec) Lower intensity = closed stomata Lower intensity & closed stomata = lower assimilation Lower assimilation & closed stomata = higher internal CO Time of experiment (min) 45 6 ambient CO 2 4 Increase Assimilation (umol/m2 sec) 4 2 Assimilation respiration Internal CO internal CO2 (ppm) Higher intensity = higher assimilation Higher assimilation & closed stomata = lower internal CO 2 Lower internal CO 2 = opening of stomata Time of experiment (min) Trigger by Light Different Speed than by CO 2 So Light Flecks are not especially useful, if high light is of short duration. Windows in canopies must give enough duration of light to allow stomata response (3+ minutes) 8
9 Leaf area Development of a plastochron index that can be used to describe the physiological age of each leaf Based on leaf area (or even leaf length) Development of an easy-to-use program for field measurement Allows one to standardize measurements based on leaf age critical to reduce data variability and to understand how flush growth influences whole tree physiology Plastochron Index application The amount of carbohydrate as related to Hass leaf age perseitol mannoheptulose sucrose Day zero = 5% of maximum leaf size 9
10 Stomata (regulated opening in leaf) CO 2 Transport from air to inside leaf - H 2 O Transport from inside leaf to air CO 2 Assimilation [Photosynthesis] H 2 O Loss From Leaf Loss of Leaf Carbohydrate / Oil Production Transpiration H 2 O Atmospheric Relative Humidity H 2 O Leaf Water Potential Xylem Conductance Stomatal Conductance Water Flow Within The Plant H 2 O Soil Water Potential Root Conductance 1
11 1 8 Fuerte Hass 2 Assimilation (μmol / m sec) Note that assimilation is falling because lower water potential induces more closed stomata and so lower conduction of both water and CO 2 flow Pre-dawn Leaf Water Potential (in MPa) [water in soil] California afternoon: higher temperature and lower relative humidity if stomata are open, higher water loss from leaf Can the soil provide the water? AM PM conductance Time 11
12 California afternoon: higher temperature and lower relative humidity if stomata are open, higher water loss from leaf Can the soil provide the water? AM PM conductance conductance Δg s + - Δg s Time AM noon PM California afternoon: higher temperature and lower relative humidity if stomata are open, higher water loss from leaf Can the soil provide the water? AM PM conductance conductance Δg s + - Δg s Time AM noon PM Conductance Difference low in AM higher in PM null set-point high in AM lower in PM Conductance (g S ) in AM (morning) Conductance Difference = g S (in PM) g S (in AM) 12
13 Δg s Mid-Day Depression of Conductance data w/nc data w/chg RH (PM) = 45% RH (PM) = 2% Stomatal Conductance in the Morning RH (AM) = 45% Measurements made with LiCor Porometer on one leaf at a time. Conclusion: Lowering of conductance in PM made worse by low RH But only if conductance in AM is high. Assimilation Dependence Upon Light Assimilation is Light-dependent CO 2 fixation Assimilation A = ½ A max A max Light Intensity This I = K I A A = max I [ Km + I ] 13
14 Assimilation Dependence Upon Light Assimilation is Light-dependent CO 2 fixation Assimilation A = ½ A max A max Light Intensity This I = K I A A = max I [ Km + I ] Maximum Assimilation (umol/m2 sec) A max Average Stomata Conductance (mmol/m2 sec) Light Constant (μmol/m2 sec) K I Average Stomata Conductance (mmol/m2 sec) g S For Hass Avocado What do we know? Light governs assimilation but.. Stomata govern (in part) assimilation too Assimilation reduces internal CO 2 but... That low CO 2 opens stomata Water loss due to transpiration changes stomata but.. Transpiration is less for high relative humidity Soil water can mitigate this leaf water loss Can we use these facts for a model that depends only upon the environment? 14
15 Model Development: Expansion Assimilation depends upon light That changes internal CO 2 That alters the conductance That changes Transpiration which is dependent upon vapor pressure deficit (which is dependent upon air temperature and relative humidity) That changes leaf water potential (which feedback upon conductance) All in the house that Jack built! Light (I) Relative Humidity Air Temperature Leaf Water Potential Vapor Pressure Deficit (VPD) Assimilation Internal CO 2 g S (i) Transpiration Sap Flow g S ( i -1) VPD = VP leaf -VP air A = A max I K I + I A = g S (Co Ci ) Ts = g S VPD We still need soil water FINALLY Internal CO 2 Controls Conductance [CO 2 ] in A stomata Difference in CO 2 = [CO 2 ] outside [CO 2 ] inside Bulk Air Boundary Air [CO 2 ] in [H 2 O ] in [H 2 O ] out Internal Air leaf Movement of CO 2 = {conductance} x {Difference in CO 2 } Movement of CO 2 limits assimilation 3 Assimilation A max Difference in CO2 (ppm) 2 1 Air CO 2 A = ½ A max -1 This [CO 2 ] = K m Internal CO Assimilation (umol/m2 sec) = A 15
16 Validation of model: How do you measure transpiration continuously? Validation of model: How do you measure transpiration continuously? SAP FLOW (continuous heat with insulated, non-invasive probes) Three Zutano Trees (A, B, C) in Green House Combination of Environmental Conditions 3/3/ dco2 = 125 calc A B C Flow of water 1.5 Comparison between the Model and the Actual Sap Flow. Three trees (zutano) were used in the green house to monitor sap flow (transpiration) and environmental parameters were also monitored. These values were used to calculate the transpiration and that was expanded to sap flow by the known of total leaf area on the branch Time of Day (hr) 16
17 Sap Flow Measurement Branch Heat carried by sap flow = Hs Heat upwards =Hu Heat in = Hi Heat downwards = Hd Heat external out =Ho Heat Balance Hi = Hu + Hd + Ho + Hs transpiration current in insulation B heater C A stem T B = 29 o T heater = 31 o T A = 29 o thermocouple voltage out Tc B Tc C Tc A T ambient = 27 o Sap Flow T A = 3 o T B = 28 o Tc B Tc C Tc A distance up the stem 17
18 Hass on Variety of Root Stocks (Duke 7 and Toro Canyon) Young trees trimmed to only several branches; all open. Data from Claudia Fassio, Summer 26 5 Total Sap Flow (g) data fit Constant Std Err of Y Est 15.8 R Squared.9769 No. of Observations 3 X Coefficient(s).938 Std Err of Coef Weight Loss (g) Other Problems: Unequal illumination of leaves on branch being tested Age Differences of Leaves on branch Field Sap Flow Measurements July 26 rooted rooted clonal rooted pollinizer clonal rooted clonal clonal road Downhill Aspect Figure 1 N road Yellow blocks = Hass Kindly provided by ACW 18
19 Rooted vs Clonal (Duke 7) (in units of cm/sec) Clonal Hass Trees Conductance in field Rooted Comparison of the Calculated Conductance between Rooted and Clonal Rootstock in the Field. Sap flow (g/hr) converted by leaf area (m2) into transpiration (mmol/m 2 sec). Using VPD from relative humidity and air temperature, the transpiration is converted into conductance. These are total conductance, not those from LICOR measurements (which misses the boundary layer). Data are from six trees planted at a field plot at ACW, Fallbrook, spaced by 1 feet from 7/6 to 7/17/6. To obtain standard conductance multiply these numbers by 25 to obtain conductance as mmol/m 2 sec. 19
20 Rooted trees maintain morning sap flow Afternoon / Morning Ratio of Sap Flow - PM / AM average 74% 18% 1.5 std. dev. 22% 24%.21 t-test.4% average PM / AM clonal rooted rooted / clon 7/6 13.2% 15.3% /7 61.5% 16.2% /8 54.4% 86.% /9 53.2% 84.1% /1 84.6% 19.9% 1.3 Hass Trees Sap Flow clonal rooted Time of Day 7/ % 84.6% / % 76.8% /14 6.7% 111.4% / % 122.7% / % 149.2% / % 11.7% 1.57 Morning = 7AM 1AM Afternoon = 3PM 6PM Clonal compared with rooted higher peak afternoon fall-off Less total water movement in rooted = less total conductance in rooted There are posters on much of this work During the poster session, a small tree with sap flow monitor is present We three thank you for your continued support! 2
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