EEB 328F: Physiological Ecology

THE ENERGY BALANCE OF ORGANISMS IN THE FIELD Introduction: The energy budget of an organism is a description of the energy flow to and from an its surface. An example would be the energy exchange between a leaf and its surroundings. This exchange of energy, in addition to any stored or metabolic energy, determines the objects temperature. Temperature is one of the important parameters affecting an organisms performance because it directly affects the rate of most metabolic processes, and can, in the extreme, inhibit all living functions. In addition, temperature directly affects the driving force for evaporation and therefore exerts significant control over the water status of plants and animals. In this exercise, we will focus the discussion on the energy balance of a flat, simple leaf, which is the simplest example for a living organisms. Similar principles apply for animals, but the situation is more complex due to their complex shape, thickness and metabolic rate. What determines the temperature of a leaf? It is incorrect to assume that temperature of a leaf is equal to air temperature. Usually it isn't, even at night. Three factors play the major role in determining leaf temperature. They are 1) the net rate of radiative heat exchange between a leaf and its surroundings, 2) the net rate of convective heat exchange between the leaf and its surroundings, and 3) the net rate of latent heat exchange between the leaf and its surroundings. Radiative heat exchange refers to energy transfer between an object and its surroundings in the form of electromagnetic (photon) radiation. Convective heat exchange arises from energy transfer as a result of collisions between the molecules in a solid (in our case the molecules making up the leaf surface) and a fluid (for example, air molecules). In the case of leaves, latent heat exchange refers to energy transfer due to evaporation or condensation of water. A fourth variable, the storage of energy, is important in thick tissue (> 5 mm), such as stems of cacti, and bodies of water. In leaves, however, storage of heat is usually negligible and is ignored. Metabolic energy is important in animals, but only rarely in plants (one plant example where heat is important is in the spadix of voodoo lilies, where heat production by high rates of respiration drives off volatile scents which attract pollinators). In a leaf, there is rarely enough metabolic activity to raise its temperature. (By contrast, in endothermic animals, metabolic energy is a major term in the energy balance equation). If a leaf is at constant temperature, the sum of the radiative (RN), convective (H) and latent (L) heat exchange processes is zero, or 0 = RN + H + L If the sum is positive the leaf temperature is increasing; if the sum is negative, the temperature is decreasing because heat is being lost to the surroundings. Radiative Heat Exchange: The net rate of radiative heat exchange is the sum of all radiation striking a leaf minus the radiation emitted from the leaf in the form of infra-red radiation. Radiation striking the leaf comes in the following forms:
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1. Short wave radiation: This is composed of radiation emitted from the sun or artificial lights and is generally considered to be of wavelengths less than 1000 nm. (Longer wavelengths are absorbed by the atmosphere). Solar radiation is made up of direct beam, diffuse beam, and reflected radiation. Photons that travel straight from the sun to the leaf make up the direct beam component. Photons that are scattered by the atmosphere or objects around the plant make up the diffuse beam, and photons that reflect off of the soil and objects below the leaf comprise the reflected radiation. On a clear day, most of the short-wave radiation is made up of direct beam photons (about 600 to 800 watts per square meter). The diffuse beam and reflected beam components on a clear day may range from 100 to 200 watts m-2. On a cloudy day, total shortwave radiation is reduced by 60% to 90%, and the diffuse component may contribute the most photons. In summary Stotal = Sdirect + Sdiffuse + Sreflected where S stands for short wave radiation. The final value of incoming short-wave radiation is modulated by a) the angle of the leaf relative to the angle of direct beam solar radiation, and b) the absorbtance of the leaf in the visible and near infrared region of the spectrum. 2. Long-wave radiation: All objects in the universe radiate energy. The amount of radiation emitted is equal to the emissivity of an object times the Stephan-Boltzman constant (; equal to 5.67 x 10-8 Watts m-2 K-4) times the absolute temperature of the object raised to the fourth power, or Remitted = - eT4 The emissivity, e, is a reflection of the molecular properties of a substance and is wavelength dependent. A black body has an emissivity of 1, a perfectly reflecting white body in the IR region of the spectrum has an emissivity of 0. Leaves typically have an emissivity in the infrared region of 0.94 to 0.98. For objects below 100oC, the radiation emitted is relatively low energy wavelengths in the infra-red region. A square meter of your naked body emits about 480 Watts to the surroundings, assuming you have an emissivity of 0.95. With clothes on, it is less (why?). In the field all objects around a leaf emit radiation according to their temperature. The sky is also an emitter of radiation, but far less than objects on the ground because of the low density of the atmosphere. When we refer to the temperature of the sky, we are referring to temperature required to emit a given amount of radiation. Typically, a clear sky has a temperature of about -20oC. How much radiation does the sky emit? The universe has a background temperature of 4oK. What does this refer to? The total radiation received by the leaf equals Rin = IRsurroundings + Stotal where IRsurroundings is the infra-red radiation absorbed from the leaf's surroundings. IRsurroundings is a composite of the IR emitted by the matter surrounding the leaf. It equals the IR incident on the
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leaf times the absorbtivity of the leaf in the far infrared portion of the spectrum, which is nearly equal to leaf emissivity. Just as the surroundings emit infra-red radiation as a function of temperature to the fourth power, the leaf also emits radiation to the surroundings. This represents the only form of radiative heat loss from the leaf. Thus, Rout = - 2eleafT4 We multiply by two because we are dealing with both sides of a leaf. The total net radiation exchange between the leaf and the environment equals RN = Rin + Rout When RN is positive, the leaf absorbs energy via radiative heat transfer. When RN is negative, the leaf gives off energy to its surroundings via radiative heat transfer. On clear nights, RN is usually negative, because the sky acts as a huge radiation sink, receiving far more radiation then it returns. On cloudy nights, the added water vapor in the clouds reradiates energy back to the ground and so radiative heat loss is reduced. Effective sky temperature on cloudy nights may be near 0oC. This explains why clear nights are associated with the coldest mornings. Convective heat transfer Convective heat transfer is the transfer of energy associated with kinetic interactions between leaf and air molecules. Essentially, convection occurs when air molecules in the boundary layer collide with molecules at the surface of the leaf and transfer kinetic energy. This changes the kinetic energy of those air molecules in the innermost boundary layer and they are replaced by other air molecules from the outer boundary layer. The more energetic molecules always transfer energy to the less energetic molecules. Hence heat transfer is always from the warmer substance to the cooler. If the air is warmer then the leaf, convection acts to warm the leaf, while if the leaf is warmer then the air, convection acts to cool the leaf. The rate of convective heat transfer depends upon the thickness of the boundary layer and the temperature gradient between leaf and air as follows: H = -2Kair (Tleaf - Tair)/B where B is boundary layer and Kair is thermal conductivity of air (0.026 watts m-1 C-1 at 25C). The boundary layer thickness depends on the size of the leaf (D) in the direction parallel to the wind speed, and is inversely dependent on wind velocity (V) D B (mm) = 4 mm s-1/2(---)1/2 V
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Convective heat transfer slows as boundary layers thicken because the molecular motion of molecules is reduced in the boundary layer, also termed the still-air layer. Wind reduces the boundary layer and more rapidly cools an object warmer than air because molecular mixing, and hence convection is increased. Wide objects have thicker boundary layers than narrow objects, and therefore, exchange energy with the surroundings more slowly. (Wind chill reflects the effect of wind on the boundary layer of mammals. Since we are warmer than the air on cold days, wind removes heat from exposed skin faster then still air does. So a wind chill of -10oC removes heat from the skin at the same rate as air would on a day where there was no wind and the air temperature was -10oC. If the air was 40oC, one may see a wind-warm effect if wind is present. Blow-dryers warming cold toes are a good example of the wind warm effect, where the rate of convective heat input is elevated by the fan in the dryer). Latent Heat Transfer Because water has such a high heat of vaporization (the amount of energy required to convert water from the liquid to the vapor state), it effectively cools materials when it evaporates. You know this already from the cooling effect associated with evaporating sweat. The heat of vaporization of water is 44,000 joules/mol. To calculate the amount of energy removed from a surface by evaporation, you only need to know the evaporation rate, that is, in the case of plants, the transpiration rate. In leaves, latent heat transfer equals L = - 2(44,000 joules/mol) x Transpiration Although transpiration in well-stirred cuvettes can be directly measured by porometers, in the field, there is no convenient way to measure transpiration of leaves without severely altering the environment around the leaf. Therefore, we need to estimate the individual parameters affecting transpiration and calculate transpiration based on those parameters. Transpiration equals Transpiration (E) = gleaf(Wleaf - Wair) where gleaf is leaf conductance and Wleaf and Wair are the absolute water concentrations (in mole fraction, i.e. mol H2O per mol air) in the leaf and air, respectively. To calculate mole fraction, we simply need to divide the vapor pressure by the atmospheric pressure. In the leaf, the vapor pressure equals the saturation vapor pressure at the leaf temperature. These values are readily available in tables such as the one you are provided with. The vapor pressure in the air equals the saturation vapor at the air temperature times the corresponding relative humidity. Relative humidity is the absolute vapor pressure in air divided by the saturation vapor pressure corresponding to the temperature of the air. Leaf conductance is the inverse of leaf resistance. We work with conductances because they are directly proportional to vapor flux. However, conductances in series are not additive. Therefore, we often utilize resistances, because in series, resistances are additive. In leaves the major resistances to water vapor diffusion out of the leaf are the stomata and the boundary layer. Total leaf resistance therefore equals
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rleaf = rstomata + rbl where rbl is the boundary layer resistance. Stomatal resistances can be determined with a porometer; boundary layer resistances must be estimated using empirically derived formula. The boundary layer resistance is a function of the thickness of the boundary layer, and therefore is dependent on wind, leaf size and leaf shape. Equipment for Measuring Energy Budgets 1. Temperature (using thermocouples and an infra-red thermometer): Thermocouples work on the Peltier principle, where at the junction of two dissimilar metals, an electrical potential is created that is proportional to temperature. Electrical potentials are small, typically around 0.05 millivolt per degree and require expensive microvoltmeters to resolve differences of a few degrees. However, thermocouples can be made quite small, allowing for temperature measurements of very small points. They are inexpensive, easily made, highly accurate and stable. Thermocouples require a reference voltage to which they can be compared. This can be supplied by simply measuring the difference in voltage between thermocouples touching the item under scrutiny and those in contact with a known standard, such as an icebath, or a copper block whose temperature has been determined by some other device, such as a thermistor. The infrared thermometer measures the IR coming into the sensing head of the instrument, and using the Stephan-Boltzmann equation which is programmed into the instrument's software, reads out the temperature of the object being measured. The instrument requires you to input emissivity, which is 0.97 for leaves. 2. Humidity (using a sling-psychrometer): This technique uses a wet bulb and dry bulb thermometer to determine relative humidity. The rate of cooling of the wet bulb due to latent heat loss is dependent on the relative humidity. Psychometric tables are usually used to relate dry and wet bulb temperatures to relative humidity. This technique is most useful from 20% to 80% RH. To use, simply wet the wet bulb with deionized water and swirl in the air. Quickly read the temperatures of the two thermometers and look up the %RH in the tables. 3. Radiation Sensors A. Li-Cor Pyrenometer: The Li-Cor pyrenometer determines the energy content of shortwave solar radiation (300 to 1000 nm) by means of a photovoltaic sensor. Output is in watts m-2. It resembles the Li-Cor quantum sensor and can be used with identical output meters as quantum sensors, given that the proper calibration constant is used. However, it should not be used under artificial lights as it has been calibrated for the daylight spectrum.

B. Fritschen-type radiometer: This device employs a thermopile, (a bundle of thermocouples) to measure the temperature of a black surface. The temperature of the surface will be a direct reflection of incoming radiation if no convective or latent heat exchange are allowed. The hemispherical polyethylene domes over the black surface ensure this. The radiometer measures radiation of all wavelengths. Originally, the radiometer we use was set up to act as a net radiometer, with an exposed thermopile on both sides of the instrument. The difference in temperature between top and bottom was a direct reflection of the difference in radiation coming in from above and below. I have modified this radiometer to yield total radiation measurements. This was done by placing an aluminum dome painted black on the inside on the bottom of the radiometer in place of a polyethylene dome. Then, I connected a thermocouple to the aluminum dome. Because aluminum has a high heat transfer capacity, its temperature is uniform and equal to that of the thermocouple. If you know the temperature of the dome, and its emissivity is 1.0 (because of the black paint), then you know the radiation absorbed on the bottom of the radiometer by the Stephan-Boltzmann law. The radiometer will still read the difference in temperature between the top and bottom surfaces, but because we know the radiation hitting the bottom, and we know the net value between top and bottom, we can subtract to get the amount of radiation coming in from the top. 4. Dataloggers: We will use a Li-Cor 1000 datalogger, Veriteq microdataloggers and/or a Campbell Scientific CR-21 datalogger to record the output from the thermocouples and radiation sensors. These instruments are programmed to give readout in degrees and radiation units instead of volts or amps. 5. Wind anemometer: An anemometer measures wind speed. We will use a hot-wire anemometer to measure wind speed. This instrument heats a fine wire to near 800 oC. The current needed to maintain the wire at that temperature depends on the rate of sensible heat loss, which is dependent on wind speed. The electronic program in the instrument converts the current to a m s-1 output. 6. Li-Cor 1600 porometer: This instrument measures transpiration of leaves in a well-stirred chamber and the temperature of leaf and air in the chamber. Hence, it is used to measure stomatal conductance. Careless people use the porometer to measure transpiration in the field directly. This cannot be done because the fan changes the environment around the leaf from that of the natural environment. The principle of operation is as follows. A leaf is enclosed in the chamber, and the fan activated. The relative humidity in the chamber is electronically measured and rises as the leaf transpires. A pump moves the chamber air through a desiccant and back to the chamber. The desiccant removes water vapor, offsetting the effect of transpiration. Ideally, the researcher adjusts the flow so that the rate of drying of the air due to the desiccant exactly offsets the rate of transpiration such that humidity is constant. References 1. Ehleringer, J.R. 1989. Temperature and Energy Measurements. Chapter 7 in Plant Physiological Ecology. Field methods and Instrumentation. RW Pearcy, JR Ehleringer, HA Mooney, PW Rundel, eds, Chapman and Hall, New York. pp. 117-136. 2. Pearcy, R.W. 1989. Measurement of transpiration and leaf conductance. Chapter 8 in Plant Physiological Ecology. Field Methods and Instrumentation. RW Pearcy, JR Ehleringer, HA Mooney, PW Rundel, eds, Chapman and Hall, New York. pp. 117-136.
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