Physiological Influences on Carbon Isotope Discrimination in Huon Pine (Lagarostrobos Franklinii)

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Physiological Influences on Carbon Isotope Discrimination in Huon Pine (Lagarostrobos Franklinii) Oecologia (Berlin) (1985) 66 :211-218 Physiological influenceson carbon isotope discrimination in huon pine (Lagarostrobos franklinii) R.J. Franceyl, R.M. Gifford2, T.D. Sharkey3*, and B. Weir3 I CSIRO Division of AtmosphericResearch, Private Bag No. 1, Mordialloc, Victoria, 3195,Australia 2 CSIRO Division of Plant Industry, P.O. Box 1600, Canberra City, A.C.T. 2601, Australia 3 Department of EnvironmentalBiology RSBS,Australian National University,P.O. Box 475, CanberraCity, A.C.T. 2601,Australia Summary. Measurements of the light environment and sto- plants were accurately predictable, then the d13C value of matal and photosynthetic behaviour are reported for Huon tree-rings from long tree ring records would provide infor- Pine (Lagarostrobos franklinii, family Podocarpaceae) in mation on past atmosphericdi3C values. western Tasmanian rainforest. For a variety of microenvi- Penget al. (1983)give an excellentsummary of the con- ronments, these are used in an analysis of stable carbon troversy surrounding the global carbon budget (and in par- isotope measurements in the air, and in branch and leaf ticular the CO, arising from fossil fuel combustion versus material, using a model for carbon isotope fractionation forest removal), and of the attempts to seeka solution using in leaves(Farquhar etal.1982). tree ring d13Crecords. The major features of d13C variations with respectto A serious diffrculty with the deconvolution of tree ring branch position can be explained in terms of the direct d13C variations to give atmosphericvariations is evident, influence of light level acting via the rate of CO, assimila- for example,in a comparison of the tree ring d13C results tion. In addition a relatively constant d13C gradient of of Freyer (1979)and Francey (1981).Each describescoher- aboat 2.61o between leaf tip and branch wood is observed. ent d13Ctrends over the last 1-2 centuriesusing many trees Alternative explanations are advanced for the tip-to- from their respective regions. Freyer's trees from Europe wood gradient in d13C. If the d13C of leaf tips is taken and USA indicate a large consistentd13C decline of almost to represent the value for photosynthate, maintenance res- 2/oobetween 1850 and 7975 A.D., while Francey's results piration is proposed as a mechanism for the further frac- from Tasmania indicate a practically constant d13C over tionation; a significant 13C depletion in respired CO, is the same period (a decline of more than about 0.21, is implied which is not supported by indirect measurements excluded). of atmospheric isotope ratio. Furthermore, an assumption Inherent in most published deconvolutions of atmo- of significant sampling errors (e.g. related to humidity ef- sphericCO, from tree rings (e.g.Stuiver 7978,Freyer7979, fects on assimilation) is required to obtain good quantita- Peng et al. 1983)is an assumption that on average,fraction- tive prediction of the light influence. ation by plant processesis constant.Tans and Mook (1980) If the branch wood d13C is taken to representthat of and Leavitt and Long (1983) empirically allow for "local the photosynthate, the tip-to-wood gradient may hnd an influences" by removing variance associatedwith tempera- explanation, via the model, in terms of tip tissue comprising ture and precipitation records (generally available over re- carbon from immature cells. Translocation of photosyri- cent decadesonly, and even then, typically accounting for thate from exposed to shaded branches is then proposed a small proportion of the variance). In addition, there is as a means of obtaining quantitative agreement with the a consistentbias towards selectionof isolated trees based predicted light infl uence. on unsupported assumptions concerning build up of re- The support provided for the applicability of the Farqu- spiredCO, around photo-synthesizingtrees in a forest. har et al. (1982)model in the field is discussedin the context Following a suggestion(Francey 1981) that variations of the problem of obtaining past global atmospheric CO, in the carbon isotope fractionation by the tree might explain levelsfrom d13Cin tree-rinss. the differences in tree-ring isotopic trends, the purpose of this work is to examinethe physiologicalaspects of carbon isotope fractionation in Huon Pine (Lagarostrobosfrank- linii; Quitn 1982).Huon Pine is a long-lived speciescon- The d13C value of a tree-ring reflects the carbon isotope f,rnedmainly to river margins in southwest Tasmania. The ratio of CO, in the atmosphereat the time the carbon longevity ofindividual trees, the ability to cross date annual was fixed, and the fractionation of CO, by plant processes rings, the frequent dominance of individual specimensin which occur between the time carbon is f,rrstfixed and when the rainforest community and the abundance of fallen and it is eventually laid down in the tree ring (Farquhar etal buried logs (which are particularly resistant to decay) con- 1982; Francey and Farquhar 1982). If fractionation by tribute to the value of Huon Pine for isotope studies over recent millenia. Offprint requestsro.' R.J. Francey The gas exchange characteristics of Huon Pine leaves * Present address: Desert Research Institute, P.O. Box 60220 were measured for different light intensities, age of shoot, Reno, Nevada 89506-0200,USA and position within the canopy and are comparedto dl3C 212 values of leavesand wood from correspondinsmicroenvi- Table 1. Average dayJong(14.2 h) photosynthetic photon flux den- ronments. The comparison is basedon the Faiquhar et al. sities (PPFD) and daily fluxes at an exposed site overlooking the (1982)model of carbon isotope discrimination during pho- StanleyRiver Valley in 1981 tosynthesis,in which dl3C of the photosynthateis relrated primarily to CrfCo, the ratio of internal to external CO, Date Average photon flux density Flux (pmol concentrations for a leaf. The measurementsform part ol m-2 s-1) (mol m-2) a major study of isotopes in tree rings, focussed on Huon 4 February 121 o 5 February 820 ia 6 February 930 49 7 February 885 40 Thus the highest hour-long PPFD which the seedling Methodology and measurements was ever likely to experiencewas about 60 pmol m-, s-1 which in plant physiologicalterms is very low indeed (com- This section briefly summarizesrelevant technical informa- 'full pared to >2,000 pmol m-2 s-1 in sun'). An average tion from Francey et al. (1984),and includesmeasurement hour-long PPFD during daylight of between i and 20 pm-ol resultsto be discussed. m-2 s-t is more likely during the summer months. Occa- sional sunfleckswere observedat both sub-canopy sites, Expedition details with the highest instantaneous PPFD neat the seedlins ob- servedat approximately 150 pmol Several expeditions have been mounted to a small area of m-2 s-1. rainforest on the Stanley River. The site is 19 km from the Tasmanian west coast in an atea largely free of human Leauesand stomata of huonpine influenceprior to 1970.Prevailing winds are south westerly The "needles" of Huon Pine and the composition of the troposphere representsthat of are thin stems completely ensheathedby overlapping air over the vast southern oceansto a large extent. Rainfall bractJike leaves.The leavesare arranged in a spiral with three is high, typically around 1,900 mm per yeat, with 190 adjacent leaves forming ap- proximately one whorl of the spiral cloudy days (> 75o/ocloud cover) and annual averagedaily around the stem. On sunshineduration of 4.8 h. each bractlike leaf, stomata were found to occur on an adaxial surface (with poor Atmospheric composition, light level and stomatal con- exposure to the open atmo- . sphere), and on the ductance measurementswere first made during February abaxial surface in two frelds divided by a ridge running length-wise 1981. Full gas exchangecharacteristics of attached shoots down the leaf. The epidermis was covered with a thick were measuredin the field in February 1982.Leaf atd wood waxJike layer, in some places completely obscuring samplesfor d13C analyseswere collected on both expedi- stomata, elsewhere only partially blocking holes over stomata. tions, as were seedlingsfor propagation and further mea- We have not been able to look at guard cellsof Huon Pine. surementsunder controlled conditions. Atmospheric com- position measurementswere continued in September1982. Slomatal conductance measurements ( February 19 8 1 ) Characteristicsof the light enuironment(February 1981) Stomatal conductance was estimated using a ventilated dif- Quantum flux density in the photosynthetically active wave- fusion porometer.Leaves were selectedfrom both seedlings band 400-700 nm, was measured on a treelessridge over- and mature trees, from low in the forest to high in the looking the StanleyRiver, and 10 minute integralsrecorded canopy, and from various degreesof shadingranging from continuously over four days. Resulting photosynthetic pho- permanently well-illuminated to permanently shady. With ton average fluxes, and flux densities (PPFD), are summa- Huon Pine it was necessaryto obtain a successionof esti- rized inTable 1. mates over 5-10 min and to exttapolate back to insertion It was raining on February 4 until mid-morning and time as the readings began to decline immediately after in- overcast all day. On February 5 7 it was sunny with scat- sertion of individual needlesinto the porometer; first read- tered cloud. This is atypical for the region which averages ings were taken within 30 s of insertion. Measured needles only 3 clear days(<25o/o cloud cover)per month over sum- were subsequentlyharvested and a micrometer used to esti- mer. mate projected leaf area. At each location light intensity Concurrent measurements were made at two forest- was determined using a pocket light meter, calibrated in floor locations,one having very little, and the other moder- sunlight againstthe quantum sensors. ate understory vegetation.At the frrst of thesesites 1.40% No clear pattern of conductances could be discerned of the (diffuse) incident photons reached the forest floor with respectto leaf position in the canopy. The entire data on the overcastday. On February 5 under bright conditions set are plotted against light intensity in Fig.
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