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This article was downloaded by: [Memorial University of Newfoundland] On: 06 October 2014, At: 15:38 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Journal of Sustainable Forestry Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/wjsf20 Nutritional Augmentation of Jeffrey Pine Saplings on a Harsh Sierran Site Roger F. Walker a a Department of Natural Resources and Environmental Science , Knudtsen Renewable Natural Resources Center, University of Nevada , Reno, Nevada, USA Published online: 24 May 2011. To cite this article: Roger F. Walker (2011) Nutritional Augmentation of Jeffrey Pine Saplings on a Harsh Sierran Site, Journal of Sustainable Forestry, 30:4, 263-283, DOI: 10.1080/10549811.2010.490107 To link to this article: http://dx.doi.org/10.1080/10549811.2010.490107 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http://www.tandfonline.com/page/terms- and-conditions

Nutritional Augmentation of Jeffrey Pine Saplings on a Harsh Sierran Site

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This article was downloaded by: [Memorial University of Newfoundland]On: 06 October 2014, At: 15:38Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registeredoffice: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK

Journal of Sustainable ForestryPublication details, including instructions for authors andsubscription information:http://www.tandfonline.com/loi/wjsf20

Nutritional Augmentation of Jeffrey PineSaplings on a Harsh Sierran SiteRoger F. Walker aa Department of Natural Resources and Environmental Science ,Knudtsen Renewable Natural Resources Center, University ofNevada , Reno, Nevada, USAPublished online: 24 May 2011.

To cite this article: Roger F. Walker (2011) Nutritional Augmentation of Jeffrey PineSaplings on a Harsh Sierran Site, Journal of Sustainable Forestry, 30:4, 263-283, DOI:10.1080/10549811.2010.490107

To link to this article: http://dx.doi.org/10.1080/10549811.2010.490107

PLEASE SCROLL DOWN FOR ARTICLE

Taylor & Francis makes every effort to ensure the accuracy of all the information (the“Content”) contained in the publications on our platform. However, Taylor & Francis,our agents, and our licensors make no representations or warranties whatsoever as tothe accuracy, completeness, or suitability for any purpose of the Content. Any opinionsand views expressed in this publication are the opinions and views of the authors,and are not the views of or endorsed by Taylor & Francis. The accuracy of the Contentshould not be relied upon and should be independently verified with primary sourcesof information. Taylor and Francis shall not be liable for any losses, actions, claims,proceedings, demands, costs, expenses, damages, and other liabilities whatsoever orhowsoever caused arising directly or indirectly in connection with, in relation to or arisingout of the use of the Content.

This article may be used for research, teaching, and private study purposes. Anysubstantial or systematic reproduction, redistribution, reselling, loan, sub-licensing,systematic supply, or distribution in any form to anyone is expressly forbidden. Terms &Conditions of access and use can be found at http://www.tandfonline.com/page/terms-and-conditions

Journal of Sustainable Forestry, 30:263–283, 2011Copyright © Taylor & Francis Group, LLCISSN: 1054-9811 print/1540-756X onlineDOI: 10.1080/10549811.2010.490107

Nutritional Augmentation of Jeffrey PineSaplings on a Harsh Sierran Site

ROGER F. WALKERDepartment of Natural Resources and Environmental Science, Knudtsen Renewable Natural

Resources Center, University of Nevada, Reno, Nevada, USA

Broadcast fertilization with an array of amendments was exam-ined for its capacity to reinvigorate growth and enhance nutritionof a 12-yr-old Jeffrey pine (Pinus jeffreyi Grev. & Balf.) plan-tation growing on an acidic Sierra Nevada surface mine site.Selected amendments consisted of Viking Brand 21-7-14, FreeFlow 29-3-4, High N 22-4-6 + Minors, and Milorganite 6-2-0 +Iron—formulations that differed substantially in critical charac-teristics including N sources and the duration of release, andeach was administered using three rates of application. All for-mulations stimulated sapling growth during some stage of thestudy, especially when applied at the highest rates, but the FreeFlow amendment, which features urea as the predominant Nsource, the High N formulation, which is a controlled releasefertilizer, and Milorganite, an organic amendment based onmunicipal biosolids, sustained growth enhancement longer thanthe Viking amendment, which relies exclusively upon ammoniacaland nitrate N forms and lacks any provision for metering nutri-ent release. As indicated by foliar analysis, increased availabilityand uptake of N probably accounted for most of the added growthinduced by fertilization, although improved P nutrition likely con-tributed as well. However, in addition to the N and P responses,fertilized saplings were frequently lower in Mn, B, and Al—all of

The author is indebted to J. Chacon, R. Fecko, G. Fernandez, W. Frederick, R. Guebard,C. McCarthy, J. Murphy, and J. Spurlock for their assistance.

Financial support for this research was provided by the Nevada Agricultural ExperimentStation and the McIntire-Stennis Cooperative Forestry Research Program.

Address correspondence to Roger F. Walker, Professor of Forest Resources, Departmentof Natural Resources and Environmental Science, Knudtsen Renewable Natural ResourcesCenter, 1000 Valley Road, University of Nevada, Reno, NV 89512, USA. E-mail: [email protected]

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264 R. F. Walker

which may be phytotoxic at elevated concentrations. Further sup-port for the possible linkages between foliar concentrations notedabove and sapling growth responses were provided by the concen-trations of these elements in the mine soil, which was low in N andP but high in Mn, B, and Al. This study reports approaches to nutri-tional augmentation on degraded sites suitable for use during thesapling stage of tree development.

KEYWORDS reforestation, forest restoration, mine reclamation,forest fertilization, forest nutrition, phytotoxicity, Jeffrey pine,Pinus jeffreyi

INTRODUCTION

Although many forest soils in temperate zones are infertile—a limitationon productivity that may involve one or several essential elements—thosedeposited as spoil materials following surface mining operations are par-ticularly prone to nutrient deficiencies, most typically of nitrogen andphosphorus (Binkley, 1986; Fisher & Binkley, 2000). Because unaidedrevegetation of mine soils through primary succession is often deemedunacceptable due to the extended time lapse between disturbance and arecovery sufficient to restore some semblance of on-site productivity andminimize off-site environmental perturbations, vegetative covers are fre-quently planted in intensive efforts to stabilize such sites and reestablishnative plant communities (Brown, Amacher, Mueggler, & Kotuby-Amacher,2003). If these efforts involve trees, however, reports of diminished sur-vival on both routine (Greaves, 1978; Powers & Ferrell, 1996; Roth &Newton, 1996) and surface mine (Czapowskyj, 1973; Vogel, 1981; Walker,West, McLaughlin, & Amundsen, 1989) reforestation sites if nutrient defi-ciencies are addressed at the time of planting, especially with conventionalsoil amendments, have fostered a reluctance to fertilize concurrently withplanting. One obvious solution for this problem is to delay fertilizer applica-tion until stands are reliably established, including a delay until the saplingstage of tree development when the greatest threat of mortality has passed,which also offers the ancillary advantage of avoiding waste associated withfertilization of seedlings that fail to survive. Furthermore, the sapling stageencompasses a period of typically accelerated biomass accrual but also onein which pronounced stunting can occur if nutrient availability is inadequateto support rapid growth, a malady demonstrated to be of particular concernin mine soils (Singh, Jha, & Singh, 2000).

Any proposal to administer nutrient amendments to a mine soil for pur-poses of stimulating sapling growth invokes two overriding decisions, themost immediate of which is the type of formulation to apply. Borrowing

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Nutritional Augmentation of Jeffrey Pine Saplings 265

from the extensive review by Tisdale, Nelson, and Beaton (1985), mostreadily available are conventional water soluble fertilizers, which are gener-ally inexpensive and capable of promptly inducing a growth response, butnutrient delivery, and thus stimulatory effects, may be relatively temporary.Controlled-release formulations greatly extend nutrient delivery, and poten-tially by extension the resulting growth response, but relatively few choicesin composition are available and their costs are generally higher. Organicamendments are available in even more limited variety than controlled-release formulations, their suitability for use in mine soils is contingent uponwhether the mineralization rates necessary for adequate nutrient release aremaintained, and adjustments in application approach to accommodate theirconsiderable mass are unavoidable, but they can provide the additionaladvantage of acting as a physical soil conditioner which may be of par-ticular benefit to mine soils given their typical paucity of organic matter(Katzur & Haubold-Rosar, 1996; Fisher & Binkley, 2000; Lunt & Hedger,2003). Subsequent to decisions regarding amendment formulation are thoseof applications rate. Because of a scarcity of documented trials at the saplinggrowth stage on either routine or degraded sites, rates at present must bederived in part from those used in trials with seedlings, which have beenincreasingly documented in recent years (Carlson, 1981; Carlson & Preisig,1981; Arnott & Burdett, 1988; Powers & Ferrell, 1996; Roth & Newton, 1996;Paquin, Margolis, & Doucet, 1998; van den Driessche, 1999; Walker, 1999a,1999b, 2002a, 2002b, 2005; Singh, Jha, & Singh, 2000; Nilsen, 2001; Vejre,Ingerslev, & Raulund-Rasmussen, 2001; Fan, Moore, Shafii, & Osborne, 2002;Clemente et al., 2004). Along with an upward adjustment to reflect the sizedifference between the two developmental stages, however, other consid-erations in adjusting these rates for use with saplings include the necessityof applying amendments at or very near the soil surface to minimize dis-turbance of established root systems, which reduces nutrient accessibilityand increases inadvertent waste, as several of the seedling trials citedabove (Carlson, 1981; Carlson & Preisig, 1981; Walker, 1999a, 1999b, 2002a,2002b; Nilsen, 2001) involved fertilization at planting with amendment place-ment directly in the root zone and thus in very small amounts. A final,and obviously crucial, consideration in selecting an application rate is thecomposition of the chosen amendment formulation, particularly regardingconcentrations of the nutrients deemed most critical for the site in question.

Reported here are the results from an investigation of the growth andnutritional responses of Jeffrey pine saplings on an eastern Sierra Nevadasurface mine to fertilization with an assortment of amendments—includingconventional, controlled release, and organic formulations—each applied atmultiple rates reflecting the chemical composition of the individual fertiliz-ers. Foliar analysis permitted a physiological interpretation of the responsesto treatment regarding alleviation of nutritional deficiencies and potentialphytotoxicities.

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266 R. F. Walker

MATERIALS AND METHODS

Study Site

An inactive, open-pit sulfur mine at an elevation of 2200 m in the easternSierra Nevada provided the study site (38◦42′30′′N, 119◦39′15′′W). Excavationceased in 1962, while active overburden placement outside the pit pro-duced a mine complex of approximately 100 ha. Spoil materials are derivedfrom hydrothermally altered volcanic rock, mostly andesites, and the minesoil is predominantly porous silica with small amounts of montmorilloniteclays (Butterfield & Tueller, 1980). The average annual precipitation in theimmediate vicinity of the mine is 50 cm and consists primarily of snow-fall. Undisturbed forest stands adjacent to the mine complex indicate thatJeffrey pine was predominant at the study site prior to excavation withCalifornia white fir (Abies concolor var. lowiana [Gord.] Lemm.) and Sierralodgepole pine (Pinus contorta var. murrayana [Grev. & Balf.] Engelm.)of lesser prevalence. At present, the vegetative cover is sparse and con-sists primarily of Jeffrey pine established through natural recolonization nearthe mine periphery and scattered plantings that created small plantations ofvarying age.

A Jeffrey pine plantation established 12 yr prior to this investiga-tion on a level spoil bench of approximately 0.5 ha was chosen forthe study. Originally planted on a 3 m × 3 m square spacing usingcontainerized seedlings, the average spacing between saplings at studyinstallation was approximately 3.5 m due to scattered mortality duringthe intervening years. To characterize the mine soil prior to treatment,five soil subsamples were collected at a depth of 0 to 30 cm from eachcorner and from the center of the bench and combined into one com-posite sample per location for a total of five composite samples. Thecomposite samples were air dried for 30 days, sieved to pass a No.10 (2.0-mm opening) screen, and analyzed as follows: texture by thehydrometer method; organic matter by loss on ignition; pH by glass elec-trode on a 1:1 mixture (by weight) of soil and distilled water; total Nby macro-Kjeldahl digestion; P (Bray 1) colorimetrically after extractionwith NH4F and HCl; K, Ca, Mg, and S by inductively coupled plasma(ICP) spectroscopy after extraction with NH4C2H3O2; Fe, Mn, Zn, Cu, andB by ICP spectroscopy after extraction with HCl; and Al by ICP spectroscopyafter extraction with KCl (Page, Miller, & Keeney, 1982; Klute, 1986). Theseanalyses revealed the following physical and chemical properties: 67% sand,19% silt, and 14% clay (sandy loam textural class); organic matter, 0.1%; pH,4.5; total N, 638 µg g−1; P (Bray 1), 23 µg g−1; K, 260 µg g−1; Ca, 3596µg g−1; Mg, 383 µg g−1; S, 294 µg g−1; Fe, 303 µg g−1; Mn, 101 µg g−1;Zn, 3.7 µg g−1; Cu, 23.7 µg g−1; B, 1.0 µg g−1; and Al, 215 µg g−1. Thecoarse texture and near absence of organic matter in this mine soil, cou-pled with the scarcity of growing season precipitation, suggest that moisture

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Nutritional Augmentation of Jeffrey Pine Saplings 267

stress imposes a limitation on growth sufficient to compromise the success ofnutritional augmentation. Furthermore, compared to routine eastern Sierranforest soils occupied by Jeffrey pine (Johnson, Susfalk, & Dahlgren, 1997;Walker, 1999a; Murphy, Johnson, Miller, Walker, & Blank, 2006), it is moreacidic and lower in N and P but higher in all of the other elements includedin the analyses, and the disparities are sufficiently pronounced for bothnutritional deficiencies and phytotoxicities to be of concern in reforestationefforts.

Study Installation

For the study, 65 test saplings were selected from among those growingon the bench, and other than a stipulation that each one be ≥ 6.0 m fromall others selected and the exclusion of stems exhibiting forking or windbreakage, test saplings were chosen randomly. One of 13 treatments wasthen assigned to each of five stems randomly selected from among testsaplings, creating a completely randomized experimental design. The treat-ments consisted of administering one of four amendment formulations usingone of three application rates per formulation plus a nonfertilized control.The formulations were: (a) Viking Brand 21-7-14 Royale fertilizer (HydroAgri North America, Inc., Tampa, FL, USA); (b) Free Flow 29-3-4 Poly-Sfertilizer (Free Flow Fertilizer, Maumee, OH, USA); (c) High N 22-4-6 +Minors controlled release fertilizer (Scotts Company, Marysville, OH, USA);and (d) Milorganite Greens Grade 6-2-0 + Iron organic fertilizer (MilwaukeeMetropolitan Sewerage District, Milwaukee, WI, USA). Amounts of individualnutrients supplied by each formulation appear in Table 1, with N indicatedby source. Application rates for the first three formulations were 250 g, 500 g,and 750 g per sapling; while those for the latter formulation were 1000 g,2000 g, and 3000 g. The fertilizers were applied in October by surface broad-casting, without tillage into the soil, in a 1.0-m-diameter circle centered atthe sapling base.

Growth Measurements

Initial measurements of sapling height and stem diameter at the ground linewere made at fertilization, with subsequent measurements completed 1 yrlater at the conclusion of the first posttreatment growing season and thenagain at the conclusions of the third and fifth growing seasons thereafter.These dimension measurements were used to calculate an estimate of shootvolume by the formula of Ruehle, Marx, and Muse (1984). For dimensionmeasurements and volume estimates, relative growth was calculated afterthe first posttreatment season based on sapling size at fertilization (seasons0–1), after the third posttreatment season based on sapling size at the con-clusion of the first posttreatment season (seasons 1–3), and after the fifth

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268 R. F. Walker

TABLE 1 Percent by Weight of Macronutrients and Micronutrients Provided byViking Brand 21-7-14, Free Flow 29-3-4, High N 22-4-6 + Minors, and Milorganite6-2-0 Fertilizer Formulations

Fertilizer

Viking Brand Free Flow High N MilorganiteNutrient 21-7-14 29-3-4 22-4-6 6-2-0

N (ammoniacal) 11.0 1.2 5.9 0N (nitrate) 10.0 0 5.3 0N (urea) 0 27.8 10.8 0N (total) 21.0 29.0 22.0 6.0a

P (P2O5) 7 3 4 2.0K (K2O) 14 4 6 0Ca 0 0 1 0Mg 0 0 1 0S 5.0 3.9 3.0 0Fe 0 2 1 4.0Mn 0 0 0.1 0Zn 0 0 0.05 0Cu 0 0 0.05 0B 0 0 0.02 0Mo 0 0 0.001 0

Note. aDerived from municipal biosolids.

season based on the size at the end of the third season (seasons 3–5). Byaccommodating differences in initial size, these relative growth calculationsprovide a more accurate assessment of early, intermediate, and long-termgrowth responses to treatment, respectively.

Foliar Analysis

Current-year needle samples were collected from the upper one-third crownof every test sapling during the 4th week of July in the first-, third-, andfifth-posttreatment growing seasons. The needles were approximately 80%elongated when collected. Segregated by sapling source and time of collec-tion, all samples were dried at 75◦C for 24 hr, ground to pass a 20-mesh(850-µm opening) screen, and then analyzed for total N using a Leco ModelFP428 N Analyzer (Leco Corp., St. Joseph, MI, USA); and for P, K, Ca, Mg, S,Fe, Mn, Zn, Cu, B, and Al by ICP spectroscopy after wet ashing with HNO3

and HClO4 (Helrich, 1990).

Statistical Analysis

Initial dimensions and volume, relative growth calculations, and nutritionaldata derived from this completely randomized experiment were subjectedto one-way analysis of variance (ANOVA) with five replications of eachof 13 treatments, and treatment effects were considered significant only

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Nutritional Augmentation of Jeffrey Pine Saplings 269

when p ≤ .05 according to the F test. Differences among means wereevaluated using Duncan’s New Multiple Range Test (DNMRT) with α =.05. All statistical analyses were accomplished using the Statistical AnalysisSystem (SAS Institute, Inc., Cary, NC, USA). In the presentation of resultsthat follows, p values are included in the text when a treatment effectproved significant as determined through ANOVA, while the mean separa-tion analysis embodied in the DNMRT was considered supplementary in thisregard.

RESULTS

Growth Responses

No influence of pending treatment on the initial height, diameter, or vol-ume of the test saplings was revealed by ANOVA, and no differences amongthe treatments prior to their implementation were revealed for any of thesevariables by the DNMRT (Table 2). Both statistical tests also indicated that,regardless of variable, relative growth during the first posttreatment growingseason was unaffected by treatment. Thereafter, however, responses to fer-tilization became readily apparent in all three growth variables. For height,relative growth for seasons 1–3 was greater (p = .0021) in saplings that hadreceived any application rate of the Viking or Free Flow formulations orHigh N at the high rate than that displayed by nonfertilized saplings; andfor seasons 3–5, it was greater (p = .0109) in those fertilized with the FreeFlow or High N amendments regardless of application rate, with the Vikingformulation at either the low or high rates, or with Milorganite at eitherthe medium or high rates. There was also evidence of an application rateinfluence on height growth within individual formulations, most apparentlywithin the Viking and High N treatments for seasons 1–3 and again withinthe latter for seasons 3–5. In each of these cases, stimulation by the highrate exceeded that by the low rate and with the stimulation by the mediumapplication interposed between them.

Relative diameter growth for seasons 1–3 was greater (p = .0022) insaplings fertilized with any rate of the Viking or Free Flow amendments,with the high rate of High N, or with Milorganite at either the mediumor high rates than that in the control treatment (Table 2). For seasons 3–5,however, diameter growth exceeding that of the control (p = .0348) occurredonly in saplings that had received any application of High N, the high ratesof the Free Flow or Milorganite formulations, or the low rate of the latter.Evidence of an application rate influence within formulations on diametergrowth was most apparent within the High N treatments for seasons 1–3and within those of the Free Flow amendment for seasons 3–5. However,in the case of the former, stimulation by the high rate exceeded that by the

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270

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Nutritional Augmentation of Jeffrey Pine Saplings 271

low rate; while in the latter, the disparity occurred between the high andmedium rates.

For relative volume growth, all fertilized saplings except those that hadreceived the High N or Milorganite amendments at the low rate or the for-mer at the medium rate exhibited higher relative increases for seasons 1–3(p = .0007) than that of the control (Table 2). For seasons 3–5, however,only the high rates of the Free Flow, High N, and Milorganite formulationsproduced such a disparity relative to the control (p = .0425). Nevertheless,some influence of application rate was evident within formulations for vol-ume growth, most apparently for seasons 1–3 within the High N treatmentsand then within the Milorganite treatments for seasons 3–5. For the former,stimulation by the high rate exceeded that by the low rate; while for thelatter, stimulation by the high rate exceeded that by either of the other rates.

Nutrition

Among macronutrients, ANOVA revealed that foliar N (p < .0001), P (p =.0488), K (p = .0360), and Ca (p = .0487) were influenced by fertility treat-ment during the first posttreatment growing season (Table 3). For N, theViking and Free Flow formulations, irrespective of application rate, and themedium rate of the Milorganite amendment produced higher concentrationsthan that in the control treatment. Within the Viking and Free Flow formu-lations, DNMRT also indicated substantial application rate influences, withthe concentrations associated with the high rates exceeding those producedby the low rates for both of these formulations and exceeding that pro-duced by the medium rate for the latter. Comparatively, P responses wereless pronounced, with the totality of the significant differences amountingto higher concentrations induced by the high rate of the Free Flow andHigh N formulations than by all other treatments except for the medium andhigh Milorganite applications. Foliar K was unique among the macronutri-ents affected by treatment during the first season in that fertilization did notelevate its concentration above that in the control according to DNMRT, andperhaps the most apparent treatment influence was one of application ratewithin the Free Flow treatments with the high rate producing a higher con-centration than the low rate. For Ca, clear evidence of fertilization resultingin a higher concentration than that in the control treatment was limited tothe low rate of the Free Flow formulation; but among the significant dispar-ities in foliar Ca, this treatment also had a higher concentration than thoseassociated with the high rates of the other three amendments.

Treatment effects on micronutrient concentrations were limited to B(p = .0134) during the first season, with the concentration in the controltreatment exceeding those of all others except for the high rate of theFree Flow formulation and the low and medium rates of the High N and

Dow

nloa

ded

by [

Mem

oria

l Uni

vers

ity o

f N

ewfo

undl

and]

at 1

5:38

06

Oct

ober

201

4

TAB

LE3

Folia

rConce

ntrat

ions

of

Nutrie

nts

and

Alin

Jeffre

yPin

eSa

plin

gsD

uring

the

Firs

tG

row

ing

Seas

on

asA

ffec

ted

by

Conve

ntio

nal

,Controlle

dRel

ease

,an

dO

rgan

icFe

rtili

zer

Form

ula

tionsa

Mac

ronutrie

ntco

nce

ntrat

ion

(%)

Mic

ronutrie

ntco

nce

ntrat

ion

(µg

g−1)

Form

ula

tion

and

applic

atio

nra

teN

PK

Ca

Mg

SFe

Mn

Zn

Cu

BA

l(µ

gg−

1)

Vik

ing

21-7

-14

250

g1.

34cd

e0.

20b

0.91

cd0.

23ab

c0.

09a

0.06

a31

a48

1a28

a8.

8a36

cd19

8ab

500

g1.

47bcd

0.19

b0.

95bcd

0.21

bc

0.09

a0.

05a

24a

498a

37a

8.0a

36cd

180a

bc

750

g1.

66b

0.20

b1.

03ab

cd0.

22bc

0.09

a0.

05a

35a

564a

33a

8.4a

38bcd

142c

Free

Flow

29-3

-425

0g

1.44

bcd

e0.

19b

0.88

d0.

29a

0.11

a0.

06a

48a

593a

37a

7.8a

36cd

188a

bc

500

g1.

63bc

0.19

b0.

97ab

cd0.

23ab

c0.

11a

0.05

a44

a56

1a38

a8.

2a38

bcd

181a

bc

750

g1.

96a

0.25

a1.

12a

0.24

abc

0.09

a0.

07a

47a

482a

38a

9.4a

40ab

cd15

9bc

Hig

hN

22-4

-625

0g

0.96

g0.

19b

0.98

abcd

0.25

abc

0.10

a0.

05a

32a

295a

35a

7.2a

48ab

232a

500

g1.

15ef

g0.

20b

1.00

abcd

0.24

abc

0.09

a0.

06a

35a

582a

34a

7.0a

41ab

cd17

3abc

750

g1.

14ef

g0.

24a

1.07

ab0.

20c

0.11

a0.

07a

23a

526a

34a

7.0a

36cd

164b

cM

ilorg

anite

6-2-

010

00g

1.02

fg0.

19b

0.93

bcd

0.23

abc

0.11

a0.

06a

25a

381a

38a

7.6a

42ab

cd23

3a20

00g

1.30

def

0.22

ab1.

08ab

0.28

ab0.

11a

0.08

a28

a88

9a40

a7.

0a45

abc

231a

3000

g1.

13ef

g0.

21ab

1.05

abc

0.21

bc

0.10

a0.

08a

32a

700a

38a

6.8a

32d

200a

bN

onfe

rtili

zed

0.96

g0.

19b

0.98

abcd

0.22

bc

0.10

a0.

06a

30a

637a

41a

9.4a

49a

240a

Not

e.aW

ithin

each

elem

ent,

mea

ns

shar

ing

aco

mm

on

letter

do

not

diffe

rsi

gnifi

cantly

atα

=.0

5ac

cord

ing

toD

unca

n’s

New

Multi

ple

Ran

geTe

st;

n=

5fo

rea

chco

mbin

atio

noffo

rmula

tion

and

applic

atio

nra

te.

272

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Oct

ober

201

4

Nutritional Augmentation of Jeffrey Pine Saplings 273

Milorganite amendments (Table 3). There was also evidence of an applica-tion rate influence on B within the High N and Milorganite treatments, withthe concentration associated with the low rate exceeding that associatedwith the high rate within the former and the concentration associated withthe medium application higher than that of the high application within thelatter. Foliar Al responses to treatment (p = .0426) were similar to those of Bin that the concentration in the control was highest overall, with the DNMRTidentifying significant differences between the control and the Viking, FreeFlow, and High N formulations applied at the high rate among other dispar-ities. Here also was evidence of an application rate effect, specifically withinthe Viking and High N formulations, as Al concentrations associated withthe low rate exceeded those at the high rate for each of these amendments.

During the third growing season, N (p = .0069), P (p = .0414), andCa (p = .0468) were the macronutrients influenced by treatment (Table 4).For N, the Viking, Free Flow, and High N amendments, regardless of theamounts supplied, along with Milorganite applied at the high rate producedconcentrations exceeding that of the control. There was also an apparentapplication rate effect within the High N and Milorganite treatments, with thehigh rates inducing higher foliar N than the low rates. For P, all rates of theViking and Milorganite amendments, along with the Free Flow and High Nformulations supplied at the high rate, produced concentrations exceedingthat in the control; and an application rate influence was evident in thelatter two formulations, as well as with the high rate inducing higher Pthan either of the other rates within both. Evidence of fertilization increasingfoliar Ca during the third season was totally lacking, as the concentration inthe control was the equivalent of the highest of those found in any of theother treatments and was greater than those associated with the high ratesof every formulation along with the medium application of the Free Flowamendment.

Among micronutrient concentrations during the third season, those ofMn (p = .0419), Zn (p = .0492), and B (p = .0474) were affected by treatment(Table 4). For Mn, the concentration in the control exceeded those associ-ated with the high rate of every formulation, as well as with the mediumapplications of the Viking and High N amendments. Within the Free Flowtreatments, an application rate influence was evident as well, with higherfoliar Mn associated with the low than with the high application. The Znconcentration in the control did not differ significantly from that of anyother treatment, and perhaps the most apparent influence on Zn was one ofapplication rate within the Free Flow treatments where the low rate resultedin a higher concentration than the high rate. Some evidence of a fertilizationeffect on B was provided by the higher concentration in the control than thatassociated with the low rate of the Viking formulation, the low and high ratesof the High N formulation, and the high rate of Milorganite. A somewhatmore apparent influence of fertilization was that on Al, as its concentration

Dow

nloa

ded

by [

Mem

oria

l Uni

vers

ity o

f N

ewfo

undl

and]

at 1

5:38

06

Oct

ober

201

4

TAB

LE4

Folia

rConce

ntrat

ions

of

Nutrie

nts

and

Alin

Jeffre

yPin

eSa

plin

gsD

uring

the

Third

Gro

win

gSe

ason

asA

ffec

ted

by

Conve

ntio

nal

,Controlle

dRel

ease

,an

dO

rgan

icFe

rtili

zer

Form

ula

tionsa

Mac

ronutrie

ntco

nce

ntrat

ion

(%)

Mic

ronutrie

ntco

nce

ntrat

ion

(µg

g−1)

Form

ula

tion

and

applic

atio

nra

teN

PK

Ca

Mg

SFe

Mn

Zn

Cu

BA

l(µ

gg−

1)

Vik

ing

21-7

-14

250

g0.

96bc

0.09

ab0.

55a

0.27

a0.

09a

0.14

a62

a75

9abc

31bc

2.4a

35b

362a

500

g0.

97bc

0.09

ab0.

48a

0.23

ab0.

09a

0.10

a49

a69

3bc

40ab

c2.

0a38

ab24

0abc

750

g0.

95bc

0.09

ab0.

51a

0.20

b0.

08a

0.09

a53

a67

4bc

36ab

c2.

0a38

ab22

8cFr

eeFl

ow

29-3

-425

0g

0.96

bc

0.08

bc

0.49

a0.

27a

0.10

a0.

11a

54a

869a

b48

ab1.

8a39

ab27

2ab

500

g1.

06ab

c0.

08bc

0.47

a0.

20b

0.09

a0.

13a

75a

801a

bc

38ab

c2.

0a42

ab23

8bc

750

g1.

06ab

c0.

10a

0.46

a0.

20b

0.08

a0.

12a

70a

491c

27c

3.4a

40ab

201c

Hig

hN

22-4

-625

0g

0.99

bc

0.07

c0.

48a

0.25

ab0.

09a

0.10

a71

a72

6abc

33ab

c2.

8a35

b29

2ab

500

g1.

02ab

c0.

08bc

0.47

a0.

25ab

0.08

a0.

10a

84a

683b

c31

bc

1.8a

43ab

228c

750

g1.

23a

0.11

a0.

58a

0.20

b0.

09a

0.12

a59

a43

1c43

abc

2.2a

36b

211c

Milo

rgan

ite6-

2-0

1000

g0.

88cd

0.09

ab0.

57a

0.27

a0.

11a

0.13

a69

a93

4ab

53a

2.2a

44ab

296a

b20

00g

0.93

bcd

0.09

ab0.

49a

0.27

a0.

08a

0.15

a59

a89

1ab

38ab

c1.

4a49

ab29

9ab

3000

g1.

14ab

0.09

ab0.

53a

0.21

b0.

08a

0.19

a70

a62

2bc

36ab

c1.

2a33

b28

9ab

Nonfe

rtili

zed

0.83

d0.

06c

0.53

a0.

27a

0.09

a0.

13a

69a

1135

a42

abc

1.8a

56a

383a

Not

e.aW

ithin

each

elem

ent,

mea

ns

shar

ing

aco

mm

on

letter

do

not

diffe

rsi

gnifi

cantly

atα

=.0

5ac

cord

ing

toD

unca

n’s

New

Multi

ple

Ran

geTe

st;

n=

5fo

rea

chco

mbin

atio

noffo

rmula

tion

and

applic

atio

nra

te.

274

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nloa

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oria

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ober

201

4

Nutritional Augmentation of Jeffrey Pine Saplings 275

in the control was greater than that associated with the high applications ofall formulations except Milorganite, as well as with the medium applicationsof the Free Flow and High N amendments. Furthermore, an application rateinfluence on Al was evident within all but the Milorganite treatments, withhigher concentrations associated with the low than with the high applica-tions for the Viking, Free Flow, and High N amendments—a result that alsoextended to the medium application for the latter formulation.

The only macronutrient affected by treatment during the fifth growingseason was N (p = .0497), which was higher in saplings that had receivedthe Free Flow or Milorganite amendments at the high rates than in thecontrol (Table 5). Among micronutrients, however, treatment influences onZn (p = .0150) and B (p = .0276) concentrations during the final seasonwere revealed. Perhaps the more prominent of several significant differencesamong treatments for Zn was a higher concentration in the control than thatassociated with the low rate of the Viking formulation and a higher onein saplings that had received the medium rate of Free Flow than in thosethat received the high rate. For B, the only significant differences were ahigher concentration in the control than those associated with the high ratesof either the High N or Milorganite amendments. A treatment influence onfoliar Al (p = .0455) was also revealed by ANOVA, but significant differencesamong treatments were limited to a higher concentration in the control thanthat associated with the high rate of the High N formulation.

DISCUSSION

The fertilizer formulations selected for trial in this study were chosenbecause they represent commonly available soil amendments and becauseof the pronounced differences among them in certain potentially key prop-erties. In particular, the four formulations differ in the N forms they featureand the extent to which nutrient release is prolonged, with the latter fac-tor dependent in part upon the former with respect to critical N nutrition.Borrowing from Tisdale, Nelson, and Beaton (1985), ammoniacal and nitrateN, which constitute the two forms in the Viking 21-7-14 formulation andserve as two of the three sources in High N 22-4-6, are water soluble andimmediately plant available, but the latter is especially susceptible to lossthrough leaching. Comparatively, urea, which supplies nearly all of the N inthe Free Flow 29-3-4 formulation and comprises the largest source in HighN, is also water soluble but must undergo transformation to provide plantavailable ions and is subject to volatilization losses with surface applications.The municipal biosolid N source in Milorganite 6-2-0 must undergo trans-formation to become plant available as well, but it is largely water insolubleand both leaching and volatilization losses are negligible. As for the durationof release, the Viking fertilizer has no provision for metering the discharge of

Dow

nloa

ded

by [

Mem

oria

l Uni

vers

ity o

f N

ewfo

undl

and]

at 1

5:38

06

Oct

ober

201

4

TAB

LE5

Folia

rConce

ntrat

ions

of

Nutrie

nts

and

Alin

Jeffre

yPin

eSa

plin

gsD

uring

the

Fifth

Gro

win

gSe

ason

asA

ffec

ted

by

Conve

ntio

nal

,Controlle

dRel

ease

,an

dO

rgan

icFe

rtili

zer

Form

ula

tionsa

Mac

ronutrie

ntco

nce

ntrat

ion

(%)

Mic

ronutrie

ntco

nce

ntrat

ion

(µg

g−1)

Form

ula

tion

and

applic

atio

nra

teN

PK

Ca

Mg

SFe

Mn

Zn

Cu

BA

l(µ

gg−

1)

Vik

ing

21-7

-14

250

g0.

82ab

0.16

a0.

79a

0.23

a0.

10a

0.11

a51

a53

0a31

c3.

2a31

ab23

7ab

500

g0.

83ab

0.20

a0.

95a

0.23

a0.

13a

0.12

a41

a63

7a46

abc

4.8a

38ab

257a

b75

0g

0.89

ab0.

19a

0.92

a0.

24a

0.12

a0.

13a

44a

759a

42ab

c5.

8a45

ab28

7ab

Free

Flow

29-3

-425

0g

0.82

ab0.

19a

0.85

a0.

22a

0.12

a0.

12a

47a

571a

42ab

c5.

6a31

ab22

5ab

500

g0.

87ab

0.22

a0.

97a

0.25

a0.

13a

0.15

a64

a90

3a57

a3.

6a45

ab27

9ab

750

g0.

97a

0.21

a0.

94a

0.21

a0.

11a

0.13

a43

a47

8a38

bc

7.2a

30ab

220a

bH

igh

N22

-4-6

250

g0.

83ab

0.19

a0.

88a

0.24

a0.

12a

0.10

a45

a31

1a46

abc

7.6a

33ab

259a

b50

0g

0.87

ab0.

18a

0.83

a0.

20a

0.10

a0.

09a

45a

492a

44ab

c4.

2a32

ab20

8ab

750

g0.

89ab

0.22

a0.

95a

0.20

a0.

12a

0.13

a49

a48

8a48

ab3.

8a24

b17

5bM

ilorg

anite

6-2-

010

00g

0.86

ab0.

22a

0.94

a0.

28a

0.14

a0.

13a

69a

530a

57a

5.8a

42ab

256a

b20

00g

0.92

ab0.

16a

0.81

a0.

28a

0.12

a0.

16a

50a

974a

46ab

c1.

6a44

ab27

9ab

3000

g0.

99a

0.21

a0.

92a

0.20

a0.

11a

0.15

a52

a63

7a51

ab5.

8a25

b24

3ab

Nonfe

rtili

zed

0.77

b0.

17a

0.90

a0.

26a

0.13

a0.

16a

65a

747a

53ab

4.6a

49a

311a

Not

e.aW

ithin

each

elem

ent,

mea

ns

shar

ing

aco

mm

on

letter

do

not

diffe

rsi

gnifi

cantly

atα

=.0

5ac

cord

ing

toD

unca

n’s

New

Multi

ple

Ran

geTe

st;

n=

5fo

rea

chco

mbin

atio

noffo

rmula

tion

and

applic

atio

nra

te.

276

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ded

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Mem

oria

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ewfo

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and]

at 1

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Oct

ober

201

4

Nutritional Augmentation of Jeffrey Pine Saplings 277

any nutrient it supplies. However, High N dispenses its contents for at leastthree growing seasons with subsurface application in eastern Sierra Nevadasoils (Walker, 2002c) due to resin coating of the prills; while the urea in theFree Flow and High N amendments, plus polymer and S coating of one-thirdof that in the former, also provides some degree of sustained N release asdoes the organic form in Milorganite. For both urea and organic N sources,however, the rate of the transformations needed to obtain plant available N,and thus by extension the duration of release, can vary substantially amongsoils.

Based on the relative growth measures computed in this study, amend-ment formulation ultimately exerted some degree of influence in theresponses to fertilization, although perhaps less than the supplementingof scarce soil nutrients in and of itself. The quantity of fertilizer adminis-tered was an obvious determinant of the strength of the growth responsesto nutritional augmentation as well. Using relative volume increases as anindicator of overall sapling growth, the early response was negligible inso-much as neither a significant treatment effect nor any significant differencesamong treatments were detected at the end of the first posttreatment sea-son, suggesting that the immediate plant availability of N embodied in theViking formulation was largely inconsequential over the short term. Duringthe second and third seasons, however, stimulation by the nutritional sup-plements was apparent to some extent regardless of formulation, as at leastone application rate of every amendment produced greater volume growthafter three seasons relative to sapling sizes at the end of the first seasonthan that exhibited by the control. Prominent among these responses werethose to the Viking and Free Flow amendments for which all three ratesproduced greater growth than the control by a margin of at least 107% andby as much as 204%, with the larger disparities associated with the FreeFlow formulation and with the high applications. Among inferences thatcan be drawn from this result is that the two water soluble fertilizers weresomewhat more stimulatory near the midpoint of the study than the con-trolled release or organic amendments, but also that the conversion of theurea in the Free Flow amendment into plant available N apparently pro-ceeded at an acceptable pace. During the final two seasons, all except theViking formulation induced greater volume growth than the control as indi-cated by sapling sizes at the end of the fifth posttreatment season relativeto that at the conclusion of the third, but in each case significant differencesoccurred only with the high application rate. Nevertheless, the disparitiesranged from 113% with High N to 125% with Milorganite. Inferences drawnfrom these findings include an apparent diminishment of the stimulatoryeffect of the Viking formulation during the final two seasons, likely reflect-ing that its immediately available N forms contribute to their relatively rapiddepletion, the controlled nutrient release by High N probably served to sus-tain its influence on sapling growth, and the conversion of the organic N

Dow

nloa

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Mem

oria

l Uni

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at 1

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Oct

ober

201

4

278 R. F. Walker

in Milorganite to plant available forms was sufficient for this formulationto prove at least comparable to the Free Flow and High N amendmentsin growth enhancement. Additionally, however, these findings indicate thatthe growth gains from fertilization were a function of the amount applied,perhaps more so than amendment composition. Nevertheless, missing inthese results was a clear indication that of the three formulations found tostimulate growth during the last two seasons, any one of them was notablysuperior, which is somewhat of a departure from the results of an earlierstudy on post-planting fertilization of Jeffrey pine seedlings (Walker, 2005)where a controlled release amendment proved to be most effective overall,although the earlier study did not include an organic formulation.

Meaningful interpretation of the results of foliar analysis requires com-parison of nutrient concentrations with reference standards, and for thisstudy the most suitable are those for western yellow pine of Jones, Wolf,and Mills (1991). Based on such comparisons, foliar N was deficient hereduring the first growing season after treatment with one exception, specif-ically that in saplings fertilized with the high application of the Free Flowformulation, and generally became much more so as the study progressed.Similarly, P was initially deficient and became exceedingly so in the thirdseason before recovering somewhat by the fifth season to a level compa-rable to that displayed initially. Foliar K followed a pattern somewhat likethat of P except that K was marginally excessive initially before concentra-tions declined to a deficient level followed by a recovery to concentrationsnear the reference standard. In contrast, Ca concentrations remained com-parable to the reference standard throughout the course of the study, whileMg concentrations were also relatively constant throughout the study butconsistently deficient. Low soil N and the prominent role of this nutri-ent in tree growth (Binkley, 1986; Kozlowski, Kramer, & Pallardy, 1991;Kozlowski & Pallardy, 1997; Fisher & Binkley, 2000), along with concen-trations in fertilized saplings that frequently exceeded that in the controlwith disparities that sometimes increased with application rate, suggest thatenhanced N availability and uptake was the primary contributor here tothe growth stimulation induced by the various amendments. However, therewas also evidence that the growth response to improved N nutrition wassomewhat delayed, as such foliar N disparities were evident during the firstgrowing season after treatment while the resulting growth stimulation wasnot apparent until near mid study. Nevertheless, the growth response gen-erally persisted through the end of the study despite foliar N disparities thathad become relatively subdued by the last season. Given that P was low inthis mine soil as well, elevation of its availability and uptake also probablycontributed to the faster growth generally exhibited by fertilized saplings,although disparities in foliar P between fertilized and nonfertilized saplingswere less prevalent than those regarding N and were entirely absent duringthe final season. None of the remaining macronutrients were deficient in

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the soil, so assessment of treatment influences on foliar concentrations, andattempts to tie the latter to growth responses, must be considered withinthis context. Thus, the fact that foliar K was low relative to the referencestandard only during the third growing season, coupled with the lack of sig-nificant differences among treatments in foliar levels during both the thirdand fifth seasons when treatment influences on growth were readily appar-ent, suggests that the differences among treatments in foliar K during thefirst season were largely inconsequential. Additional evidence supportingthis conclusion was that despite the supply of K in every formulation exceptMilorganite, none of them elevated foliar K above that of the control at anytime and the most apparent fertilization effect on foliar levels in the firstseason was one of application rate within the Free Flow treatments, andthis amendment supplies less K than either the Viking or High N formu-lations. Likewise, attributing any growth effects to the differences amongtreatments in foliar Ca during the first season is dubious, as only saplingsfertilized with the low rate of the Free Flow amendment had a higher con-centration than the control, yet only High N supplies Ca among the fourformulations. The lower Ca in fertilized than in nonfertilized saplings dur-ing the third season, which was consistent across all formulations at theirhigh application rates, may reflect to some degree a dilution effect result-ing from accelerated biomass production (Timmer, 1991), as similar foliarCa responses have been documented previously in Jeffrey pine fertilizationtrials (Walker, 1999b, 2002c, 2005). The consistently low foliar Mg concen-trations contrasted against its high level in the soil, but any capacity of theamendments to rectify the former, including High N which has a Mg source,was never in evidence. Because no suitable reference standard has beendocumented for S, the adequacy or inadequacy of the foliar levels foundhere is unknown, but the capacity of any of the amendments to alter theselevels was also never evident despite the S supply in all formulations exceptMilorganite.

All of the micronutrients were abundant in the soil, but again usingthe Jones, Wolf, and Mills (1991) standards for comparison purposes, foliarconcentrations of these elements varied considerably in relation to thereference values. Specifically, Fe was consistently low here in all treatmentsthroughout the study, while all Mn concentrations were at least 4× thereference value, and in the most exceptional case of the nonfertilizedsaplings during the third season, was 15× the standard. Overall, Zn wassomewhat low during the first and third seasons before rising to concentra-tions near the reference value in the fifth season, while Cu concentrationsapproximated the reference value initially before declining for the third andfinal seasons. With the exceptions of those found during the fifth seasonin saplings that had received either High N or Milorganite at the high rates,foliar B concentrations were somewhat higher than the standard value.Given the lack of significant differences among treatments at any time

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during the study, a conclusion easily drawn from the foliar Fe concentra-tions is that the source contained in all except the Viking formulation wasinconsequential in alleviating a possible Fe deficiency. In contrast, the highMn level during the third season in nonfertilized saplings relative to thoseassociated with the high application of every formulation and extending tothe medium applications of the Viking and High N amendments, in light ofthe consistently elevated foliar concentrations found throughout the study,suggests that fertilization may have alleviated Mn phytotoxicity near midstudy. This element is known to reach toxic levels in acidic mine soils(Cummins, Plass, & Gentry, 1965; Barnhisel & Massey, 1969; Berg & Vogel,1973; Fisher & Binkley, 2000), and earlier studies with Jeffrey pine seedlingsconducted in the same mine complex (Walker, 2002b, 2005) demonstratedsimilar Mn responses to amendment application. Some treatment differencesin Zn concentration were revealed during both the third and fifth seasons,but foliar Zn levels seldom approached, much less exceeded, the referencestandard; and fertilization, including that with High N which contains aZn source, did not consistently modify foliar concentrations relative to thatin the control. Consequently, these differences may have been incidentalfluctuations that had tenuous, if any, ties to growth responses. Apparently,the Cu source in High N was also largely inconsequential given the completeabsence of significant differences among treatments in foliar Cu. Researchon B toxicity in forest trees (Glaubig & Bingham, 1985; Walker, 1999b) hasnot yet provided the definitive diagnostic criteria needed for its assessment,and although the B concentrations revealed here frequently exceeded thereference standard, the disparities were not especially pronounced, so itis uncertain whether the lower concentrations frequently encountered infertilized saplings relative to that in the control were indicative of fertiliza-tion acting in an ameliorative capacity. The Jones et al. (1991) referencestandards do not include a value for Al, nor is one available elsewhere,which also renders it difficult to evaluate potential phytotoxicity associatedwith the Al concentrations encountered here. Nevertheless, Al was elevatedin this mine soil as well, and fertilized saplings frequently had lower foliarAl than nonfertilized saplings, most often when the high application ratewas administered. It is not uncommon for this metal to be implicated inthe phytotoxicities that can occur in acidic mine wastes (Cummins, Plass, &Gentry, 1965; Berg & Vogel, 1973; Butterfield & Tueller, 1980), and previousstudies with Jeffrey pine seedlings on such sites have revealed a similar Alresponse to fertilization (Walker, 2002a, 2002b, 2002c, 2005).

In summary, these results demonstrate substantial growth stimulationin a 12-yr-old Jeffrey pine plantation growing on an acidic and infertileSierran surface mine site induced by surface application of an array ofnutrient amendments. Three application rates of four formulations wereexamined, and water soluble, controlled release, and organic amendmentswere included. All four formulations reinvigorated sapling growth at some

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point in the study, especially with the highest application rates, but a for-mulation featuring only ammoniacal and nitrate forms of N did not sustaingrowth enhancement as well as those containing urea or organic N sources.Improved N nutrition, and to a lesser extent that of P, likely accountedfor much of the growth stimulation by fertilizer application, althoughamelioration of potential phytotoxicities may have contributed as well.

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