METHODS OF ESTIMATING FLUXES IN THE SWARD: RATES OF LOSS

1. rates of loss from tissue turnover measurementes

Rates of loss can be determinad on the basis of number of leaves lost and their weight, and the number can be determined either by direct observation in clover or, in ryegrass, calculated from the formula n1+al-n2 where n1 and n2 are the number of leaves per tiller determined on the bench at times 1 and 2 and al is the number of new levaes per tiller observed in the field between t1 and t2. rates of tissue loss from leaves can also be determined from the rate at which tissue senesces and its weight per unit lenght. Early mesurements were made using this approach by fulano in perennial ryegrass. In the former instance of regular observations of the chanes in area of necrosed and senescent leaf were combined with its dry weight per unit area; in the latter the accumulation of total lenght of senescent tissue over a period of time was multiplied by the weight per unit lenght of dead leaf. A senescence often occurs quickly, with a time lapse between sucessive leaves, coefficients of variation for rates of senescence based on changes in leaf lenght and weight tend to be rather high, values from two experiments being 105% and 137%. The rates exhibited a skew distribuition, so that it was importante to use transect or replicate means rather than individual values and to enrure that the mean were based on sufficient tillers to normalise the distribution.

2. DIRECT METHODS The total amount of material which dies during a given period of time can be estimated by observing any changes in the pool of standing dead material and adding to this an allowance for any material which has decomposed during that time. Alternatively, dead material may be collected at frequent intervals so that decomposition can be ignored. Two methods used in ecological studies are considered first, either of which might be used when an area cannot be visited regularly. In both methods and initial sample of the herbage is taken and separated into living and dead parts. Two further areas, the control and the threated areas, are then set aside for a later harvest. Both area are sampled at the later harvest and living and dead material is again separated. In the Wiegert and Evans (1964) methods the final observations are weight of living material (m1) and dead material (d1) on the untreated plot, and residual dead material on the treated plot (dr). In the method of Lomnicki et al. the observed quantities are m1, d1, and the amount of green material which was entered the dead pool (dm). Thus the total amount of material (D) which dies in a given period is the sum of a net change in the dead pool (d1-do) and the calculate total amount of material which has disappeared D= (d1-do) +1/2 (do+d1) Ldt

The variables t and ld are respectively time and the rate of disappearance of dead material, measured in units of weight per unit of weight per unit time and given by formula (log do 0 logc rs)/t. Clearly the same calculation can be made if the Lomnicki method is used. The choice must depend on wich of the two methods is more practical under the circumstances of the experiment. indicado separar o material morto e vivo no laboratrio. MM seco, pesado e returned to the soil surface where is protected by a nylon or stainless steel mesh. With more frequent harvesting the problem of decomposition can be avoid and the total weight of dying leaves is equal to the total collected. Fulano found that removing dead matter from a ryegrass/clover sward had no effect on the dry weight of green material which was produced. Dead sheaths could not be removed without causing damage to the grass tillers. The estimation of rates of dry matter loss from any of these methods raises further questions: how much weight been lost by the dying leaves by the time they are sampled or measured? How much of this loss represents the remobilization and reclycling of organic compounds to younger leaves, and at what stage should leaf material be weighed or measured to obtain the best estimates? Evidence on the scale of these losses is in itself conflicting. Robson and Deacon (1978), studying the changes in weight leaves on the main stems of young plants of perennial ryegrass in growth chambers concluded that 30% of the weight was lost between maximum weight at full expansion and death., Vin also foun 40%, in julho agosto perdeu 20%, 60% na primavera. mto dificil achar a maneira mais adequada para achar estas respostas. Until further experimental work indicates clearly to what extent carbon fixed in one leaf reappears in another youngher leaf it is difficult to decide what is the most appropriate practice. Certainly a low estimate (not including recycled carborn) would be most appropriate when estimates of growth are based on carbon fixation as in the study of Parsons el al 1984

RATES OF DISSAPPEARANCE OF LITTER Various workers have calculated the amount of material leaving the standing dead pool (l) in a given time from the initial (do) and the final (d1) weights of material in the pool at times to and t1 and the amount of new dead material (dn) which has entered the pool during the period in question. The equation is: L=(do+dn-d1)/(t1-to) Results showed that large amounts of material vanish in wet conditions but the dry conditions dead material may tend to accumulate The rate of disappearance of litter can also be estimated using litter bags. Bocock and gilbert emphasize that the bags should have an aperture large enough to admit earthworms and mesofauna, they recommend the use of untinted nylon harnest of 10 mm aperture. Comparaes indicam q este mtodo superestima as perdas. Outros used a stainless steel mesh plate on top of the litter instead of litter bags. Similar results were obtained by Yates

using boxes wich mesh bottoms firmly pegged on the soil, the within mesh spacing being approximaltely 2mm.

DEMOGRAPHIC ANALYSIS Data on the births and deaths of leaves and tillers can be analyzed using demographic methods such as survivorship curves, life tables and probabilities of births and deaths. A plot of n1 (the number of individuals in a cohort or population surviving to age t) against t produces a survivorship curve, which can be used to determine the median survival time or half-life of the population as a whole () Mean life expectancy (et) for any individual in the population is then given by the formula: Et=(nx+e + nx+2e + nx+3r.)/nc+0.5c Where c is the time period convered by each age class

OBSERVATIONS IN GRAZED SWARDS 1. Frequency and severity of grazing The fate of marked tillers in grazed swards is of particular interest and was already being investigated in the sixties and early seventies. The tillers can be located using transects by using a large standard quadrat sectioned or by using a metal template whose free ends act as pointers to the tillers. These permit proper recolation of the transect, co-ordinates, quadrat or template. Normalmente se marca com anis de plasticos ou fios de plsticos, mas se perde atravs de pssaros, pisoteio dos animais especialmente em altas lotaes. Cores vo de vermelho, amarelo, laranje. Grazing frequency is established by examining the tillers at frequent intervals, comparisions of different frequencies showing that this should be done every two days at high grazing pressures. At low pressures twice weekly may be sufficien, since the frequency of observation depends on the expected frequency of defoliation. The amount of leaf removed has been measured as a percentage of the total length of lamina or of lamina plus shenath wich was present before grazing with a correction factor applied for the growth taking palce between observations based on increase in length of emerging leaves on similar ungrazed tillers. Coefficients of variation for defoliation frequency on a per tiller basis ranged from 35% to 96% with mean values of 55 and 67% being obtained from different experiments. The greater variability was normally experienced at the lower stocking rates.

2. FLUXES IN GRAZED SWARDS So far consideration has been given to measurements of fluxes during periods in which the swards are not concurrently being grazed. Once the grazing animal is introduced it becomes necessary account for the herbage consumed as well as the other fluxes. The original equation deltaW=deltaG-deltaI-deltaD as before, if there of the has se parameters can be measure the fourth can be calculated. A useful and productive approach has been to study fluxes in swards maintained at a fixed height or weight by variable stocking so that changes in delta W are minimized and may be ignored. So: deltaG= deltaD+deltaL the technue allows intake to be estimated as the difference between growth and senescence. The experimenter is, however, left with a difficult problem: that of estimating deltaG in a situation in which the evidence is being, so to speak, consumed in front of his or her very eyes. Fluxes in grasss Three basic approaches have been adopted to deal with the problem of determing deltaG in grazed grass swards. The first relies on measuring growth on those tillers which escaped desfoliation during the period in question, but introduces an element of bias because ungrazed tillers represent a smaller than average section of the total population. The second approach in which growth measurements are made on tillers protected from grazing by exclosure cages, involves less frequent monitoring, but is also open to some criticism. Provide, however, that grazing is not sever enough to remove more than half of the leaf or part of some of the sheaths the practice can be very largely justified in relation to length increments. Althought is usual to measure length increments on all the leaves of the tillers under observation, increments can also be assessed as the product of the rate of leaf appearance and the length of the first undamaged leaves to emerge fully on the caged tillers. A third approach to the problem of estimating the lengths of leaves grown and eaten, which avoids use of exclosure cages, has recently been adopted by tallowin. This combines measurements of net increases or decreases in leaf length with measurements of bottom up extension growth made over periods of 2-3 days on tillers marked with Indian ink at the point of emergence of the youngest leaf, and at the lamina/sheath junction of the youngest fully emerged leaf. Conversions of length increments to equivalent increments in dry weight can of course, be made on a straight weight to length ratio provided they are based on a fully representative population of intact leaves or grazes leaf remnants. This is not the case in a dynamic situation in which leaf tips are constantly being removed, and constitutes a particular problem in species in which the lamina taper sharply towards its tip, since it cannot be assumed that the weight per unit length of the lamina is constant over its full length. have provided mathematical descriptions of the different relationships between leaf shape and weight per unit length in

leaves of different shapes wich could, in principle, be used to translate the ratio of the lengths of leaf eaten and grown into an equivalent weight ratio.

Fluxes in white clover

Because of the growth of a white clover petiole ceases after the lamina has been removed estimates of growth and intake do not require the use of exclosure cages an can be obtaine in the open sward. The appropriate tees were pushed into the soil near the marked stolon, exmina-lo aps uns dias permit ever quantas folha foram comidas, apareceram e morreram. It proved quite easy in practice to distinguish the clean cut petioles of leaves grazes by sheep from those normally older The weight of incomind, eaten and uneaten leaves were obtained from trowel samples eased out of the soil to avoid stolon breakage, and analyzed in laboratory.

Utilization Can be calculed by dividing the weight of the material eaten by the weight of material grown in the same period, and is usually expressed as a percentage. It is most conveniently calculated in grasses as the difference between growth and senescence divided by growth In concluding this section it should be stressed that, since almost any attempt to measure dry matter fluxes in grazed swards involves some element of compromise, the choice of method is up to the investigator and depends on the nature of the experiment and the facilities and personnel available. Pequenos erros no importam se os efeitos forem similares ao do experiment and valuable conclusions may be drawn as long as actual numerical values are treated with necessary caution.

COMPUTATION OF RATES OF GROSS DRY-MATTER PRODUCTION AND SENESCENCE The gross increase in dry matter over a period of time is equal to the amount of tissue which enters the living pool in that time. It can also be calculated from the change in the weight of the material in the living pool, plus the total weight of material lost from it during the same time period by death. The weight of tiscsue entering or leaving the system is the product of the entry or loss per unit and the number of units involved. Unless the change in the number of these units is considerable it is reasonable to use the arithmetic mean of the initial and final determination in this calculation. The fairly erros on crop growth rates calculated as products in this way are in accordance with expectation

A method based on demographic principles was used by for a chalk grasslands community. He established the half-life of the leaves from a probit relationship between number of survivors in different cohorts and time, and added half of the initial biomass to the observed production over the period of the half-life

CONCLUSIONS In all work involving the computation of gross rates of production the golden rule is that fuxes may only be ignored if they can be shown to be of minor significance over the period of time in question. To ignore tiller deaths or losses of dead material from the litter is tantamount to assuming that no tillers die and no litter is lost. Over relatively short periods or in specific circumstances this may well be so, but it should be never be assumed to be the case without good reason. The choice of the best method in a given situation is a matter for the individual but, because of the nature of the erros involved, rates of flux should be calculated from products rather than from differences or sums wherever possible. It is sometimes technically easier to measure one kind of flux than another it is easier to determine deltaG and deltaI for clover in continuously grazed swards than it is to continue deltaD. Choices may have to made between quicker and less accurate methods or more precise but slower methods involving additional correction factors. At all times the investigator must be on the alert for possible sources of bias: error can be measured, but bias can creep in undetected. The method choosen is of course secondary to the purpose of the exercise. Tissue turnover studies are time-consuming and, before embarking on such studies, the experimenter should be sure of their value in the context of the problem he or the wishes to address. Almost certainly the future holds as many lessons for agronomists as the past.