Growth Fluctuation of Loblolly Pine Due to Periodic Air Pollution Levels: Interaction of Rainfall and Age

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1 Disese Detection nd Losses Gowth luctution of Loblolly Pine Due to Peiodic Ai Pollution Levels: Intection of Rinfll nd Age S.. Phillips, J. M. Skelly, nd H. E. Bukht ome Gdute Resech Assistnt, Associte Pofesso in the Deptment of Plnt Pthology nd Physiology; nd Associte Pofesso in the School of oesty nd Wildlife, espectively, Vigini Polytechnic Institute nd Stte Univesity, Blcksbug, VA 461. Senio utho is cuently Specil Pojects oeste, Missoui Deptment of Consevtion, Columbi, MO 651. The uthos wish to thnk the dministtos of the Rdfod Amy Ammunition Plnt, Rdfod, Vigini, fo thei coopetion, nd the Southen oest Disese nd Insect Resech Council fo the ptil funding of this poject. Accepted fo publiction 1 Decembe ABSTRACT PHILLIPS, S.., J. M. SKELLY, nd H. E. BURKHART Gowth fluctution of loblolly pine due to peiodic i pollution levels: intection of infll nd ge. Phytopthology 67: The effect of peiodic souce of NO,SO i pollution egime on the gowth of loblolly pine ws investigted, Rdil incement gowth studies wee conducted to detemine if coeltions existed between emission levels (pedictions of poduction levels wee used s suogte) of souce nd dil incement gowth of loblolly pine. Thee stnds of loblolly pine poximl to the emission souce wee smpled by obtining bestheight (1.37 m bove the bse of the tee) incement coes fom 5 tees in the dominnt o codominnt cown clsses of ech stnd. Multiple line egession nlyses utilizing nnul dil incement gowth s the dependent vible nd the independent vibles of nnul poduction levels, totl nnul infll, nnul sesonl infll, nd ge wee used to evlute dil gowth of smple tees within ll stnds. An invese eltionship significnt t P=.1 ws demonstted between gowth nd poduction levels in two of the loblolly pine stnds. Additionl nlyses of these stnds indicted theoeticl eductions in dimete gowth within both stnds without the pesence of visible injuy. Gowth nd poduction levels wee not significntly coelted in the emining loblolly pine stnd. Additionl key wods: tee gowth loss, sulphu dioxide, nitogen oxides, Pinus ted. The inhibition of gowth in foest tees due to the pesence of sevel diffeent i pollutnts hs been demonstted (, 5, 7, 8, 9, 1). Sevel eseches hve demonstted tht gowth losses cn occu without the pesence of "visible symptoms" (5, 9, 1). The "hidden injuy" theoy of plnt dmge due to i pollution s developed by Stokls (1) in 193 consideed two points impotnt to the nlysis of gowth loss: eduction in the photosynthetic ctivity of the plnts, nd educed gowth nd/o yield without symptom development. Teshow et l. (1) conducted nnul dil gowth studies with Dougls fi (Pseudotsug menziesii Mib.) suounding souce of tmospheic fluoides. They found eduction of up to 5% in the nnul dil gowth of tees subjected to the mbient fluoide pollution. This eduction in gowth ws pesent iespective of foli (o visible) injuy. Pollnshutz (9), woking with the impct of SO, hydogen fluoide, nd mgnesite dust on the gowth of sevel foest tee species, found eduction in nnul dil gowth without the ppence of visible symptoms. Hontvedt (7) pesented the wok of Stein nd Dssle who conducted dil incement gowth studies with spuce subjected to mbient SO. They found 45%, 7%, nd 18% eduction in dimete gowth ove 9y peiod which coesponded diectly to men yely mbient SO levels of 543.4, 143., nd g/min 3, espectively, Linzon (8) studied white pines (Pinus stobus L.) subjected to mbient SO fumigtions in the foest suounding the Sudbuy Smelte distict of Ontio, Cnd, to obtin n estimte of the economic impct of mbient pollution levels. He found tht thee ws n nnul stumpge loss of $14, nd tht the mket vlue loss pe ye ws $117,. The estimtes involved only white pine which compised 7.6% of the foests in the X 16 m (3 mi ) of ffected e. Clson nd Hmme () utilized nnul dil gowth to pedict the gowth impct of fluoide i pollutioninsect intection on lodgepole pine (Pinus contot Engelm.). They detemined loss of 3.7 M 3!/hecte/y (57 bod feet/ ce/ ye) bsed upon dil gowth. Extpoltion to the entie ffected e esulted in loss estimte of. ppoximtely 38,55 m3 (8.5 million bod feet) pe ye. The U.S. Amy's Rdfod Amy Ammunition Plnt (RAAP) is n industil souce of nitogen oxides (NO.)nd SO. Stone nd Skelly (11) hve conducted nnul dil incement gowth studies t the RAAP with white' pine nd yellow popl (Liiodendon tulipfe L.). The unique histoyeof thepraap, hving thee distinct peiods of poduction (Wold W II, the Koen conflict, nd the Vietnm conflict), nd the eltionship of this peiodicity to i pollution studies ws pesented by Stone nd Skelly (11). A simple line egession nlysis ws used by the eseches to evlute the eltionship of Copyight 1977 The Ameicn Phytopthologicl Society, 334 Pilot Knob Rod, St. Pul, MN All ights eseved, nnul dil incement gowth to poduction levels (n indicto of i pollution levels). They demonstted significnt invese eltionship between nnul gowth 716

2 June 1977] PHILLIPS ET AL.: AIR POLLUTION/LOBLOLLY PINE nd nnul pollution levels fo both yellow popl nd white pine. This wok incopoting moe efined sttisticl nlysis ws the bsis fo studies t the RAAP involving sevel stnds of loblolly pine (Pinus ted L.). Loblolly pine lso ws exmined owing to its impotnce to the foest industy of Vigini nd the southesten United Sttes (6). Thee wee thee distinct objectives of these investigtions: (i) to detemine wht coeltions, if ny, existed between histoic poduction levels of the RAAP nd the nnul dil gowth of loblolly pine; (ii) to evlute wht effect tee ge nd infll hd upon coeltions of gowth nd poduction levels; nd (iii) to obtin n estimte of the gowth impct of the S NO,, i pollution egime on loblolly pine within the RAAP. MATERIALS AND METHODS 717 wee conducted fo ll thee smple stnds. A stnd nlysis lso ws conducted to obtin n estimte of the effect of pst pollution levels t the RAAP on gowth of the smple stnds. This nlysis ws conducted with those stnds which exhibited significnt invese eltionship between gowth nd poduction levels fte emoving the effect of ge nd two infll vibles. The estimte ws obtined by pedicting the gowth tes of tees within the RAAP nd tees gown in n hypothesized pollutionfee condition. The estimte of these gowth diffeences ws obtined by mens of the MLR eqution which pedicted dil incement gowth fom the independent vibles of ge, totl nnul infll, nnul sesonl infll, nd nnul poduction levels t the RAAP. Estimtes of nnul gowth diffeences within individul stnds wee Rdil incement studies wee conducted in thee plnted stnds of loblolly pine to detemine if coeltion existed between pst pollution levels t the RAAP nd the nnul dil gowth tes of tees 1. LEVELS PRUCTION,oo RADIAL AVERAGE NCREMENT I GROWTH 9W 9 X 8. t 7. subjected to these levels. All of the smpled stnds wee gowing in level e on 1.8m spcing in sndy cly soil. Loblolly pine stnd No. 1 ws locted.6 km nothest of the RAAP's min powe fcility nd ws 15 CIs, 7 y old t the time of smpling. Loblolly pine stnd No. U O3 1.3.W ws 18 y old t the time of smpling nd ws locted in the sme genel e s the site of loblolly pine stnd No. 1.7 km nothest of the min powe fcility. Loblolly pine stnd No. 3 ws lso 15 y old nd ws locted 1.4 km nothwest of the min powe fcility. No silvicultul tetments hd been conducted in these stnds. The stnds wee smpled by the method of Stone nd Skelly (11). This smpling povided the vibles of nnul dil incement gowth nd ge of the tee t the time of smpling. The nnul dil incements of the smpled tees wee detemined to the neest.1 mm fo the peiod of W. C,too > 6 ". YEA, Z W GOL Z 5o U 5. R t.o A, sow 7 'o Multiple line egession nlyses wee conducted vible nd ge, totl nnul infll, nnul sesonl X ". I by Stone nd Skelly (11). Tee ges wee obtined s.i. ~i 1. 1,65 197,oo 9 so PRODUCTION LEVELS 9.o AVERAGE RADIAL GROWTH INCREMENT _4. GRO1WTHI INCREMENT TO P z 3.E 1.: VL IVCRAGERADT o96 the RAAP s independent vibles; the ltte s deived 8. infll, nd pecent vege nnul poduction levels of conducted with dt fom the individul stnds. The fou SLR nlyses wee computed with nnul dil incement gowth s the dependent vible nd nnul poduction levels, nnul sesonl infll, totl nnul infll, nd ge septely s the independent vible. The fou MLR nlyses utilized nnul dil incement gowth s the dependent vible with the following combintions of independent vibles: ge nd poduction levels; totl nnul infll nd poduction levels; nnul sesonl infll nd poduction levels; nd ge, totl nnul infll, nnul sesonl infll, nd poduction levels. These eight line egession nlyses... 1 ooo utilizing nnul dil incement gowth s the dependent peviously mentioned. Rinfll dt obtined fom the Blcksbug, Vigini Meteologicl Sttion (3) povided the emining independent vibles of totl nnul infll nd nnul sesonl (fom I Apil though 3 Septembe of the ye) infll. ou simple line egession (SLR) nlyses nd fou multiple line egession (MLR) nlyses wee.o cc> eo 5. 4.: 4o 3.,o C[,o., ig. 13. Avege nnul dil incement gowth of 5 loblolly pine tees locted in ech of thee stnds [ig. 1), ), nd 3) epesent dt fo tee stnds 1,, nd 3, espectively ] t the U. S. Amy's Rdfod Amy Ammunition Plnt (RAAP) nd the poduction levels of the RAAP fom

3 PHYTOPATHOLOGY 718 obtined fom the yely dt of infll (nnul nd sesonl), ge, nd poduction which wee substituted in the eqution simultneously to pedict men nnul dil incement fo tees subjected to pollution. This pediction of gowth ws ccomplished fo ll yes which wee included in the egession nlysis. A pollutionfee gowth ws hypothesized by intejecting the yely dt of ge, nnul sesonl infll nd totl nnul infll in the eqution. Howeve, insted of inseting the coesponding poduction level fo tht ye, zeo level of pollution ws ssumed to pedict men nnul dil incement gowth of the smple tees unde pollutionfee condition. The men nnul dil incement gowth pedicted fo both the pollution nd pollutionfee conditions within n individul stnd wee summed ove ll yes nd comped by mens of pecentges of gowth eduction. RESULTS The nnul dil incement gowth of the smple tees within loblolly pine stnd No. 1 did not demonstte significnt line eltionship with nnul poduction levels of the RAAP (ig. l, Tble 1). A significnt positive line eltionship existed between nnul dil incement gowth nd the ge of the smple tees of this stnd. The two infll vibles wee not significntly coelted with dil incement gowth in SLR nlysis. The MLR nlyses then wee utilized to compe nnul dil gowth to nnul poduction levels of the RAAP fte fist emoving the effect of othe vibles independently o in combintion. When the effect of ge ws emoved, nnul dil incement gowth demonstted significnt invese coeltion to nnul poduction levels. A significnt line eltionship ws not demonstted between gowth nd poduction fte emoving eithe of the infll vibles. A significnt invese line eltionship ws demonstted between incement gowth nd poduction levels when ge nd the two infll vibles wee emoved fist. This eltionship hd highe vlue nd ptil coeltion coefficient thn ny of the pevious egession nlyses. In loblolly pine stnd No., ll of the independent nd [Vol. 67 MLR vibles demonstted significnt invese line eltionship between nnul dil incement gowth nd poduction levels (ig., Tble 1). The highe vlues nd coeltion coefficients wee demonstted by the egessions of poduction on gowth. The egession nlysis in which ge ws emoved nd poduction egessed on gowth lso yielded high vlues s did gowth egessed on poduction fte ll othe vibles wee emoved. This lst nlysis mintined the getest degee of coeltion nd the lgest vlue. The nnul dil incement gowth of smple tees in loblolly pine stnd No. 3 hd significnt positive line eltionship to nnul poduction levels (ig. 3, Tble 1). Age lso hd significnt positive coeltion with incement gowth. A significnt invese eltionship between totl nnul infll nd incement gowth ws found lso, but no significnt eltionship ws exhibited between incement gowth nd nnul sesonl infll. When eithe of the two infll vibles wee emoved fist nd poduction in the MLR nlysis ws comped to dil gowth, n incese in positive coeltion nd significnce ws exhibited. Howeve, when ll othe vibles wee emoved o ge lone ws emoved, the eltionship between gowth nd poduction ws nonsignificnt. The diffeences between pedicted men nnul dil incement gowth of tees subjected to pst i pollution levels of the RAAP nd tees in hypothesized pollutionfee condition povided eltive estimte fo the mount of gowth eduction in the smple stnds. Only loblolly pine stnds No. 1 nd No. demonstted the necessy significnt invese eltionship of dil incement gowth to nnul poduction levels of the RAAP needed to povide n estimte of gowth eduction. The pedictive model fo men nnul dil incement gowth in loblolly pine stnd No. 1 ws Y = X, X.541 X X4 ; whee: Y = men nnul dil incement; X, = nnul poduction levels of the RAAP; X = ge; X 3 = totl nnul infll; nd X4 = nnul sesonl infll. This eqution povided n estimted dimeteinsidebk (d. i. b.) gowth of 56.7 mm fo tees subjected to the RAAP's mbient i pollution levels nd 13. mm fo the sme TABLE 1. Multiple line egession nlysis esults fo thee loblolly pine stnds t the Rdfod Amy Ammunition Plnt, Rdfod, Vigini. Dependent vible consideed ws nnul dil incement gowth fom Loblolly stnd #1 Loblolly stnd # vlues fo vlues fo Independent Regession numbe vible 1 Avege nnul poduction **.65 Age 18.6* *e *.16 3 Totl nnul infll ** Totl sesonl infll **.17 5 Avege nnul poduction 44.6** ** Avege nnul poductionb **.53 7 Avege nnul poductionc **. 8 Avege nnul poductiond 57.88** **.393 Coelted to nnul dil incement holding ge constnt. bcoelted to nnul dil incement holding totl nnul infll constnt. 'Coelted to nnul dil incement holding nnul sesonl infll constnt. dcoelted to nnul dil incement holding ll othe vibles constnt. easteisks indicte: (*) significnt vlue t P=.5 nd (**) significnt vlue t P=.1. Loblolly stnd #3 vlues fo 1.66* ** **

4 June 1977] PHILLIPS ET AL.: AIR POLLUTION/LOBLOLLY PINE tees in n hypothesized pollutionfee condition ove 1y peiod. Thus, the estimted dimete gowth inhibition is 46.5 mm o 45% eduction in d. i. b. due to the RAAP's mbient i pollution levels, The egession eqution povided fo pedicting men nnul dil incement gowth in loblolly pine stnd No. ws: Y = X X+.164 X X4.The estimted d. i. b. gowth ove 1y peiod fo smple tees in this stnd subjected to the RAAP's i pollution levels ws 78.7 mm; tht fo pollutionfee gowth ws estimted t 14.5 mm. The pedicted gowth inhibition ws 63.8 mm o 45% eduction in d. i. b. DISCUSSION The effect of i pollution on the gowth of loblolly pine hs not been evluted pio to this study. The SLR nlysis with nnul dil incement s the dependent poduction levels s the independent vible vible nd s nnul peviously used (1 ) demonstted the eltionship between i pollution levels nd gowth of the smple tees without consideing vitions in tee ge nd infll. The MLR nlysis in which nnul dil incement ws egessed on nnul poduction levels fte emoving infll nd ge effects ws consideed the most vlid ppisl of the effects of i pollutnts upon the gowth of the smple tees. The emining thee MLR nlyses tht wee conducted llowed the two infll vibles nd the ge vible to be emoved independently befoe evluting the eltionship of i pollution on gowth. The vibility of plnt's ection to i pollution often is elted to ge. This vition in sensitivity hs been demonstted with loblolly (1) nd Vigini pine (4) seedlings subjected to fumigtions with ozone. The effect of ge s it ffects pollution sensitivity ws ppent in loblolly pine stnds No. 1 nd No. t the RAAP. In loblolly pine stnd No. 1 when nnul dil incement gowth nd nnul poduction levels wee comped in SLR nlysis the eltionship ws not significnt. When the vibility in gowth due to the influence of ge ws emoved, significnt invese eltionship ws gowth nd poduction levels. The demonstted effect of ge onbetween this eltionship hd completely disguised the significnt effect of i pollutnts on the gowth of the smple tees. Simil esults wee found with the smple tees of loblolly pine stnd No.. In this stnd, both poduction levels nd ge independently wee demonstted to hve highly significnt effect upon the gowth of the smple tees. The emovl of vition ssocited with ge within this stnd demonstted tht highe invese coeltion of dil gowth to poduction levels ws pesent thn hd been obtined with SLR nlysis. The effect of ge on i pollutiongowth eltionship in this sitution hd not completely disguised this significnt eltionship s in loblolly pine stnd No. 1, but the filue to conside this vible educed the negtive coeltion between i pollutnts nd gowth. Envionmentl fctos consideed in the nlyses of stnds t the RAAP consisted of infll vibles: totl nnul infll nd nnul sesonl infll. Unlike ge, the two infll vibles did not getly ffect the coeltion between poduction levels nd gowth when 719 thei effect ws emoved fist; i.e., the invese coeltion of poduction to incement gowth becme stonge in loblolly pine stnds No. 1 nd No.. Afte emoving the effect due to the two infll vibles nd ge, the negtive ptil coeltion coefficient between gowth in loblolly pine stnd No. 3 nd nnul poduction level ws not significnt. Simil esults concening the vibility in sensitivity of tees to i pollutnts due to envionmentl fctos wee pesented by Wilhou (13). The smple tees within loblolly pine stnds No. 1nd No. filed to exhibit foli symptoms which wee chcteistic of i pollution injuy in spite of the significnt effect of i pollutnts on gowth within the stnds. Teshow et l. (1) hd demonstted this phenomenon with Dougls fi subjected to tmospheic fluoides. Loblolly pine stnd No. 3 exhibited i pollution injuy symptoms on the mjoity of the smple tees nd ws locted 1.4 km fom the min emission souces. Loblolly pine stnds No. 1 nd No. wee locted.6 nd.7 km, espectively, fom the min emission es. Theefoe, loblolly pine stnd No. 3 ws pobbly subjected to highe pollution concenttions thn stnds No. 1 nd No. owing to the gete diffusion of the pollutnts by the time they hd eched the moe distnt stnds. Loblolly pine stnd No. 3 ws situted poximl to souce of highly concentted sulfuic cid mist. Howeve, the SLR nlysis egessing incement gowth on poduction levels indicted significnt positive influence of i pollution. The effect of ge on gowth lso ws positive nd highly significnt. When the influence of ge ws held constnt the eltionship between i pollution nd gowth becme insignificnt. The coeltion between ge nd gowth duing the peiod of ws vey high. Theefoe, the positive influence of i pollution on gowth in the SLR nlysis ws esult of filing to conside tee ge s gowth vible. Poduction levels wee not coelted significntly with the gowth of smple tees within this stnd even fte emoving the effect of ge nd infll. This phenomenon could be due to tolence of the smpled tees within the stnd to i pollution injuy. The sensitivity expessed by the othe two stnds which wee moe distnt fom the souces of i pollution nd without visible injuy comped to the foli injuy within stnd No. 3 peclude this conclusion. A moe esonble explntion of these diffeences my be tht i pollution levels ne nd within stnd No. 3 my hve been sufficiently high to inhibit gowth to mximum point whee no futhe esponse to the known fluctutions in the pollution densities bove this citicl level ws detected. The fct tht smple tees within loblolly pine stnd No. 3 demonstted foli symptoms of i pollution indicted the pesence of consistently highe levels of i pollution thn wee pesent in stnds No. 1 nd No.. The sttisticl nlysis unde such conditions would not detect this sttic influence of i pollution on gowth. These esults indicte tht loblolly pine tees plnted within the RAAP my be undegoing n inhibition of gowth which is popotionl to the RAAP's poduction levels nd which is influenced by tee ge nd infll. These esults, theefoe, confom to Stokls's (1) concept of invisible injuy. It lso is pobble tht some

5 7 PHYTOPATHOLOGY [Vol. 67 loblolly pine tees my exhibit tolence to the NOxSO 5. DONOBAUER, E Tee ing nlysis s mens of i pollution egime t the RAAP. The conclusion tht dignosis nd evlution of pollutnt effects. Abstct symptom expession is not peequisite fo gowth loss No. 991 in Abstcts of Ppes, Second Int. Cong. of plces sevee estictions upon gowth loss estimtes Plnt Pthol. 51 Sept 73, Minnepolis, MN (Am. utilizing symptoms fo dmge estimtes. It, theefoe, is Phytopthologicl Society, St. Pul, Minnesot) highly possible higlyopossilel tht gowth lo t gow loss osse in tin foests fo subjected nts m6. to HARLOW, (unpged) W. M., nd E. S. HARRAR Textbook of lowlevel nd longtem exposue to i pollutnts my be dendology. (5th ed.). McGwHill, New Yok. 51 p. occuing unnoticed nd/o unevluted. 7. HORNTVEDT, R SO injuy to foests. J. o. Utiliz. 78: LINZON, S. N Economic effects of sulphu dioxide on foest gowth. J. Ai Pollut. Contol Assoc. 1:8186. LITERATURE CITED 9. POLLANSHUTZ, J Obsevtions bout the susceptibility of vious kinds of tees with espect to 1. BERRY, C. R Age of pine seedlings with pimy emission of SO, H nd mgnesite dust. Pges needles ffects sensitivity of ozone nd sulphu dioxide. in Poc. 1st Eu. Cong. Infl. Ai Pollut. Plnts nd Phytopthology 64:79. Anim. 1968; Wgeningen, The Nethelnds. 419 p.. CARLSON, C. E., nd W. P. HAMMER Impct of 1. STOKLASA, J Die beschtdigung de vegettion fluoides nd insects on dil gowth of lodgepole pine duch uchgse und fbiksexhltionen. Ubn und ne n luminum smelte in nothwesten Montn. Schwtzenbeg Velg, Belin, und Wien. 487 p. U.S. Dep. Agic., o. Sev. (Noth Reg.) Rep STONE, L. L. nd J. M. SKELLY The gowth of two P. foest tee species djcent to peiodic souce of i 3. CROCKETT, C. W Climtologicl summies fo pollution. Phytopthology 64: selected sttions in Vigini. Bulletin 53. Wte Resouces 1. TRESHOW, M.,. K. ANDERSON, nd. HARNER. Resech Cente, V. Polytech. Inst. nd Stte Univ., Responses of Dougls fi to elevted tmospheic Blcksbug, Vigini. 168 p. fluoides. o. Sci. 13: DAVIS, D. D The influence of plnt ge on the 13. WILHOUR, R. G The influence of tempetue nd sensitivity of Vigini pine to ozone. Phytopthology eltive humidity on the esponse of white sh to ozone. 63: Phytopthology 6:579 (Abst.).