HORTSClENCE 47(7):917-926. 2012. Response of Higbbush Blueberry to Nitrogen Fertilizer during Field Establishment-II. Plant Nutrient Requirements in Relation to Nitrogen Fertilizer Supply David R. Bryla1,4 U.S. Department ofAgriculture, Agricultural Research Service, Horticultural Crops Research Unit, 3420 NW Orchard Avenue, Corvallis, OR 97330 Bernadine C. Strik2 Department of Horticulture, Oregon State University, 4017 ALS, Corvallis, OR 97331 M. Pilar Baiiados3 Departmento de Fruticultura y En%gia, Pontijicia Universidad Cat6lica de Chile, Casilla 306-22, Santiago, Chile Timothy L. Righetti University of Guam, UOG Station, Mangilao, Guam 96923 Additional index words. Vaccinium corymbosum, fertilizer rate, integrated plant nutrient management, macronutrients, micronutrients, nutrient partitioning, plant dry matter, reallocation Abstract. A study was done to determine the macro- and micronutrient requirements of young northern highbush blueberry plants (Vaccinium corymbosum L. 'BIuecrop') during the first 2 years of establishment and to examine how these requirements were affected by the amount of nitrogen (N) fertilizer applied. The plants were spaced 1.2 x 3.0 m apart and fertilized with 0, SO, or 100 kg·ha-1 of N, 3S kg.ha-1 of phosphorus (P), and 66 kg·ha- 1 of potassium (K) each spring. A light fruit crop was harvested during the second year after planting. Plants were excavated and parts sampled for complete nutrient analysis at six key stages of development, from leaf budbreak after planting to fruit harvest the next year. The concentration of several nutrients in the leaves, including N, P, calcium (Cal, sulfur (S), and manganese (M"n), increased with N fertiIizer application, whereas leaf boron (B) concentration decreased. In most cases, the concentration of nutrients was within or aboyc the range considered normal for mature blueberry plants, although leafN was below normal in plants grown without fertilizer in Year 1, and leafB was below normal in plants fertilized with SO or 100 kg·ha- 1 N in Year 2. Plants fertiIized with SO kg·ha- ' N were largest, producing 22% to 32% more dry weight (DW) the first season and 78% to 90% more DW the second ~eason than unfertilized plants or plants fertilized with 100 kg·ha- 1 N.Most DW accumulated m new shoots, leaves, and roots in both years as well as in fruit the second year. New shoot and leaf I DW was much greater each year when plants were fertilized with SO or 100 kg·ha- N, I whereas root DW was only greater at fruit harvest and only when SO kg·ha- N was ap~lied. Application of SO kg.ha- I N also increased DW of woody stems by fruit barvest, but neltber SO nor 100 kg.ha- I N had a si!!11ificant effect on crown, flower, or fruit DW. Depending on treatment, plants lost 16% t: 29% of total biomass at leaf abscission, 3% to 16% when pruned in winter, and 13% to 32% at fruit barvest. The content of most nutrients in ~be plant folIowed the same patterns of accumulation and loss as plant DW. Howeycr, unhk~ ~W, magnesium (Mg), iron (Fe), and zinc (Zn) content in new shoots and lea~:es was Similar among N treatments the first year, and N fertilizer increased Nand S content m woody ~tems mUch earlier tban it increased biomass of tbe stems. Likewise, N, P, S, and Zn content III the crown were greater at times when N fertilizer was applied, wbereas K and Ca content were sometimes lower. Overall, plants fertilized with 50 kg·ba- I N produced the most gro\~~h and, from planting to first fruit harvest, required 34.8 kg.ha- I N, 2.3 ~'ha-I P, t2~,kg.ba K. 8.4 kg'ha-1 Ca, 3.8 kg.ha- Mg S.9 kg·ba- ' S 295 g·ba-I Fe, 40 g·ba B, 23 g·ba copper (Cu), ' 1273 g·ha- I Mn, and 6S g. b~-I Zn. Thus, of the total amount offertilizer ap~lied ove~ 2 years, only 21 % of tbe N, 3% of the P, and 9% of tbe K were used by plants dunng estabhshment. Like many crops, fertilizer practices in bluebeny (Vaccinium sp.) are routinely adjusted by comparing the results of leaf nutrient analysis HORTSCIENCE VOL. 47(7) JULY 2012 at a standard time against the known optimal ran"es of leaf nutrient concentrations. E fTectiv: fertilizer management, however, also requires a good understanding of plant nutritional demands both in terms of the nutrient amount (Santos, 2011) and the timing in which each nutrient is most needed (Mattson and van Iersel, 20 II). Biomass determination through sequential plant excavation. coupled with nutrient analysis of each tissue type, is presently the most reliable way to obtain the amounts and seasonal patterns of plant nutrient uptake (Weinbaum et aI., 2001). Nutrient analysis of entire plants at multiple times during annual cycles of growth and development is difficult and expensive, and only N has been examined in detail in highbush blueberry (Bafiados et aI., 2012; Hanson and Retamales, 1992; Retamales and Hanson. 1989; Throop and Hanson, 1997). Nitrogen is the predominant nutrient applied to blueberry for successful commercial growth and production. Although the blueberry plant is relatively small and slow·growing compared with many temperate fruit tree crops, the amount of N fertilizer applied to the crop each year is comparable (Stiles and Reid, 1991). Typical rates in Orcgon, for example, average 50 to 100 kg·ha- ' ofN per year during planting establishment and 100 to 300 kg·ha-' of N per year once the field matures. Other nutrients are also applied, largely based on soil tests and general recommendations from plant and soil testing laboratories. Hart et al. (2006) developed more stringent guidelines for nutrient management of blueberry based on leaf tissue and soil analysis. However, witli-me exception of N. the defined ranges of nutrient sufficiency were based on experience and not on controllcd studies of each nutrient. Nutrient requirements in perennial fruit crops such as blueberry depend on new biomass production in vegetative and reproductive tissues, the nutrients needed for production of the new tissue, and the amount of nutrient, reallocated from existing plant tissues. Mature northern highbush blueberry plants produce most new shoots and leaves in the spring and early summer, before or during fruit development. and most new roots in early spring. before budbreak, and mid· to late summer after fruit harvest (Abbott and Gough, 1987). Most N is acquired during shoot and fruit development (Throop and Hanson, 1997). and therefore split applications of granular N fertilizer are recommended in the spring (April to June in the northem hemisphere) for blueberry (Hart et aI., 2006). Reallocation of nutrients in woody plants occurs internally. especially in early spring from storage tissues such as the crown and woody stems and in the fall from senescing leaves (Mohadjer et aI., 200 I; Rempel et aI., 2004; Strik et aI., 2(04) and externally from decomposition of plant tissues such as senesced leaves and roots and pruned wood (Strik et aI., 2006). We previously found that 50kg·ha-' N per year promoted more gro\\1h and yield than no N fertilizer during establishment of highbush blueberry, whereas rates 100 kg·ha-' N or greater were excessive and resulted in salt stress and plant mortality in the young plant· ing (Baiiados et aI., 2012). Through destructive 917 harvests and the use of depleted 15N fertilizer, we estimated that unfertilized plants gained 1.6 g/plant of N from soil sources, whereas fertilized plants accumulated 2.3 g/plant of N, on average, 60% of which was from fertilizer Nand 40% was from the soil. The objectives of the present study were to determine the requirements of N and other macro- and micronutrients in the young plants and to examine how the requirements were affected by the amount of N fertilizer applied. Plants were excavated, separated into relevant plant parts, and analyzed for nutrients on six key dates from leafbudbreak immediately after planting to the first fruit harvest the next year. one row ofl2 plants. The N fertilizer used was granular ammonium sulfate (2IN-OP-OK). Three equal applications of the fertilizer were sprinkled around the base of the plants each spring at a total rate equivalent to 0 (no additional fertilizer N), 50, or 100 kg·ha- ' N per year. A fourth set of plants was fertilized at a rate of ISO kg·ha- ' N but was not used in the present study as a result of severe problems in the treatment with salt injury and plant death ---e. o· A Materials and Methods (Bafiados et aI., 2012). The fertilizer was applied on 11 Apr., 20 May, and 27 June in 2002 (Year I) and on 24 Mar., 8 Apr., and 23 June in 2003 (Year 2). Each plant was also fertilized with 35 kg·ha- ' P and 66 kg·ha- ' K each spring. Plants were winter-pruned in Feb. 2003 and lightly cropped the second year after planting. One randomly selected plant from each treatment plot was excavated and sampled Year 1 Year2 B 3.5 N rate (N) 3.0 Year 1" Year 2· N rate (N) Year 1" NS Year 2 0.16 YxN'· 0.14 ~ 0.12 ~ YxN** Two-year-old 'Bluecrop' blueberry plants were obtained from a commercial nursery (Fall Creek Farm & Nursery, Lowell, OR) and transplanted on 27 Mar. 2002 to a field located at the North Willamette Research and Extension Center in Aurora, OR. The plants were delivered from the nursery in 4-L pots filled with a mix of peat, bark, and pumice. Both roots and the soil mix were placed in the planting hole and packed firmly in place with surrounding soil. Soil at the site was a Quantama fine loam (mixed, mesic Aquatic Haploxeralfs) with a pH 5.0 and 3.0% organic matter content. The soil was amended before planting with a 10-cm deep layer of Douglas fir (Pseudotsuga menziesii Franco) sawdust and 66 kg·ha- ' ofN [2IN-OP-OK ~hS04]. The sawdust and fertilizer was incorporated 0.2 m deep by cultivation in ~ I-m wide strips to form five 50-m-Iong rows of planting beds. Plants were spaced 1.2 m apart within rows and 3.0 m apart between rows (equal to a density of 2778 plantslha). Grass alleyways (1.1 m wide) were planted and maintained between the rows. The field was irrigated as needed (~25 to 38 mmlweek) from May to September each year using overhead sprinklers. Three N fertilizer treatments were applied to plants during the first 2 years after planting. The treatments were arranged in a randomized complete block design with three replicates per treatment. Each treatment plot consisted of z ., 2.5 0······· "Cii -' Q 2.0 .. ~ 0.10 1.5 1.0 '--'----~_ _ _.>_..1 c 1.0 ~ L-..------'----L-l0.08 D N rate (N) Year 1'· Year 2 NS 0.40 N rate (N) NS Year 1 Year 2 NS YxN NS ~ 0.8 0.35 l> YxN' 0.30 !!l. ;s:: .... '" ~ 0.25 co ~ 0.20 0.6 0.15 0.4 '-'----~--_--'-..J E 0.8 iii - 0.6 '--'-----'-----~ N rate (N) Year 1" Year 2' YxN NS YxN NS 1200 .1:'L .... ... ~ '" l> 900 rJ) ~ 0.10 F N rate (N) Year l' Year 2· !!l. ;s:: :J 0.4 600 0.2 0.0 ~---~---~.J G ~ '--'-----~---~o H 600 N rate (N) Year 1NS Year2 NS 500 YxN NS 40 300 0············· 2···· l> ~ ." 400 20 ~ 300 100 Received for publication 27 Feb. 2012. Accepted for publication 14 May 2012. We thank G. Buller, H. Rempel, and M. Resendes for technical support and acknowledge financial support from the Oregon Blueberry Commission and the Northwest Center for Small Fruits Research. Mention of a trademark, proprietary product, or vendor does not constitute a guarantee or warranty of the product by the U.S. Dept. of Agriculture or Oregon State University and does not imply its approval to the exclusion of other products or vendors that also may be suitable. I Research horticulturist. lProf;:ssor and berry crop extension specialist. 'Former PhD graduate student. Department of Horticulture, Oregon State University. 'To whom reprint requests should be addressed; e-mail david.bryla@;ars.usda.gov. !!l. CIl ~ 5- 200 o~------~----~ o 50 100 N fertilizer rate (k ·h -') 9 a '--'--------~------~~100 a 50 100 N fertilizer rate (kg·ha·') Fig. 1. Leaf (A) nitrogen (N), (B) phos horus . (F) manganese (Mn) (G) boron (Bf d (ii): (C) calcIUm (Ca), (D) magnesium (Mg), (E) sulfur (5). 'B!uecrop' blueben; during the ::d ) Iron (Fe) concentrations in response to N fertil!zer rate I~ eac spnng \~ith 0, 50, or 100 kg.ha-' N and Ithe second year after planting. The plants were fertlhzed 3 (Year 2). Results from individual d eab~es were collected 24 July 2002 (Year I) and 25 July 200_ . . an com med I' f . * .. -f nonslgmficant and significant at P <: 0 05 ana YSls 0 vanance are inset in each graph: 1'S" three replicates and error bars - . and 0.01, respectively. Each symbol represents the mean 0 represent ± I SE. firs; 918 HORTSCIENCE VOL. 47(7) JULY 2012 ~ destructively on six key dates: leaf budbreak (29 Apr. 2002), late July when leaf sampling is recommended for nutrient analysis (24 July 2002; see Hart et aI., 2006), and just before leaf abscission (29 Oct. 2002) in Year 1; and just before winter pruning (7 Feb. 2003), leaf budbreak (22 Apr. 2003), and late July when fruit were first harvested (25 July 2003) in Year 2. Each plant was dug up from the entire width of the planting bed to a depth of 0.3 m to recover as much of the root system as possible. Roots were washed after digging to remove the soil, and the plants were partitioned into new shoots, leaves, flowers and fruit (Year 2 only), l-year-old wood, 2-yearold wood (Year 2 only), the crown, and roots. Each part was then oven-dried at 70°C, and dry weights were recorded after the samples reached a constant weight. New shoots and l-year-old wood produced in 2002 were reclassified as 1- and 2-year-old wood, respectively, in Feb. 2003. All dried plant parts were ground to pass through a 40-mesh (425-llm) screen and analyzed for N using a combustion analyzer (CN2000; Leco Inc., St. Louis, MO) and for P, K, Ca, Mg, S, Fe, B, Cu, Mn, and Zn using inductively coupled plasma--optical emission spectroscopy (Optima 3000DV; Perkin Elmer, Wellesley, MA) after acid-digestion (Jones and Case, 1990). Fruit samples were analyzed for N but were unavailable for analysis of other nutrients, and root samples were not analyzed for Fe as a result of potential issues with soil contamination. Data were analyzed by analysis of variance using SAS Version 9.2 software (SAS Institute, Cary, NC) and square-root transfonned when needed as determined by the heterogeneity of variance. Any transfonned data were back-transformed for presentation. Means were separated at the 0.05 level using Duncan's new multiple range test. Results and Discussion Leaf nutrient status. The concentration of N in leaf samples taken in late July increased with N fertilizer application as has been shown by others in northern highbush (Ballinger and Kushman, 1966; Bishop et ai., 1971; Cummings, 1978; Cummings et a\., 1971) and rabbiteye (V. virga tum Ait.) (Spiers, 1983) blueberry. The leafN concentrations in this study, however, were higher than what is considered normal (1.76% to 2.00%; Hart et aI., 2006) for northern highbush blueberry in Oregon, especially during the first year after planting when levels averaged 2.82% in plants fertilized with 50 kg·ha-' Nand 3.48% in those fertilized with 100 kg·ha- ' N (Fig. lA). By the second year, leaf N concentrations were lower in N-fertilized plants but again were still higher than nonnal, a~eraging 2.23% with 50 kg·ha- ' Nand 2.57% With 100 kg·ha- ' N. Without N fertilizer, leaf N concentrations averaged only 1.42% the first year after planting but increased to 1.76% the second year. It is possible that current recommendations for sufficiency of leaf N concentration (Hart et at., 2006), generally HORTSCIENCE VOL. 47(7) JULY 2012 used for producing plants, may need to be increased for young, establishing plants. Leaf N concentration has been shown to decrease with planting age (Cummings, 1978). The concentration of several other leaf nutrients also increased significantly with N fertilizer application, including P and Ca during the first year after planting (Fig. 1B-C), Mg during the second year after planting (P < 0.10; Fig. ID), and Sand Mn during both the first and the second year after planting (Fig. IE-F). Others have found little effect of N fertilizer rate on leaf P (Ballinger and Kushman, 1966; Bishop et a!., 1971; Cummings et a!., 1971) in establishing northern highbush blueberry, although variability in response has differed with site (Cummings et aI., 1971). The effect ofN fertilization on leafCa has likewise been sho\\TI • 0 kg·ha·' -0- 50 kg·ha·' -.....- 100 kgha·' ~ :e- A 180 ~ 150 ~ ~ I I r" Nxr** ~ 120 "0 a p Year 1 NNS I I 90 I 1;; ~ o/! (5 0 .s:: 0 60 I 30 I I. a I. I a Year2 0 ~ I I r" NxP* I • I" b I I b I • I c I I / (/J 0 ~ C 120 80 40 ~ 20 0 0 / i / I / I 60 ~ "0 -0 I Pruned T** NxT** .9 E p NNS III :e- 100 ~ a B / b I b a 'E III 60 C NNS :e- 50 ~ 40 E 30 .9 '" ~ "0 C ;: e () .1 T** NS NxT I I P I • 20 10 0 0 'E 200 c. '" ~150 N** r** NxT** " :::: '" E 100 ~ "0 '0 0 a: 50 0 Apr. July Oct. Jan. Apr. JUly 2002·2003 . new shoots and leaves, (B) woody stems, (e) crown, and (D) roots of :Bluecrop· Fig. 2. Dry weIght of (A)Ii (2002) and second (2003) year after planting. The plants were fertIlIzed each blueberry dunng the rst ha- 1 • (N) and pruned 7 Feb. 2003. Results from two-way anal}sls spring. with 0, 50, or IO~ k.~e m:gi~se~ in each graph: NS, ., •• = nonsignifi~nt and significant at ofvanance [N rate (i\) t1 .. I<DLch SYmbol represents the mean of three rephcatcs; means Without p ~ 0.05 and 0.01, respecltne y. ive~ date are not significantly different at the 5% level (Duncan·s letters or with a common etter on a g new multiple range test). 919 to vary by site as well as cultivar within a region (Cwnmings et aI., 1971). Bishop et al. (1971) and Ballinger and Kushman (1966) found that Ca decreased with increasing N rates in mature 'Bluecrop' and 'Wolcott' blueberry, whereas Cummings (1978) reported little effect of N rate on leaf Ca in the first growing season and a decrease \\ith N rate in the second year. Leaf Mg was affected little by N rate in our study, as has been reported by others (Ballinger and Kushman, 1966; Bishopetal., 1971; Cummings et aI., 1971). LeaH,tn concentration increased with N fertilization. which was likely related to a decline in soil pH (Baiiados et aI., 2012). also observed by Townsend (1973) in a 6-year study in 'Blueray' blueberry. In general. P and S concentrations were lower in the N-fertilized plants in the second year than they were in the first, declining by an average of 0.4 to 0.5 mg'g-' and 0.8 to 2.6 mg.g", respectively, whereas Ca, Mg, and Mn concentrations were higher the second year after planting, increasing by an average of 1.3 to 1.4 mg·g", 1.3 to 1.6 mgK', and 468 to 564 IlgK', respectively. An increase in leaf Mg. Ca, and Mn over time has been reported by others (Cwnmings, 1978; Townsend, 1973). In our study, only leaf P and Ca concentrations were within the range each year considered optimum for blueberry (i.e., 0.10% to 0.40% for P and 0.41 % to 0.80% for Ca; Hart et aI., 2006), and this was the case whether plants were fertilized with N or not. LeafMg and Mn concentrations, on the other hand, were higher than recommended (i.e., 0.13% to 0.25% for Mg and 30 to 350 Ilg·g" for Mn; Hart et aI., 2006) the second year after planting, regardless ofN treatment, and leaf S concentrations were higher than recommended (0.11% to 0.16%; Hart et aI., 2006) in both years but only when N fertilizer was applied [as (NH 4hS04]. Leaf B concentration was also significantly affected by N fertilizer application, but in this case, the response differed between years (P < 0.05; Fig. IG). In the first year after planting, leaf B concentrations increased from 48 Ilg·g" without N fertilizer to 62Ilg'g-' with 50 kg·ha" Nand 59 Ilg'g" with 100 kg·ha" N. The next year, the opposite was true, and leaf B decreased from in blueberry has been previously reported (Cummings, 1978). Most DW accumulated in new shoots leaves, and roots in both years (Fig. 2) a~ well as in fruit the second year (Table I). The total of these parts combined represented 76% to 80% of the total DW by the end of the first season and 71% to 75% by fruit harvest of the second season. New shoot and leaf DW was much higher each year with N fertilizer (Fig. 2A), whereas root DW was only higher at harvest and only when 50 kg·ha" N was applied (Fig. 2D). Application of 50 kg·ha-' N also increased DW of woody stems by fruit harvest (Fig. 2B), but neither 50 nor 100 kg·ha" N had any significant effect on crown (Fig. 2C) or flower and fruit DW (Table I). Not surprisingly, unfertilized plants allocated relatively more biomass to roots and less biomass to shoots than fertilized plants, but only during the first growing season. By the end of the first year, the rootto-shoot ratio in unfertilized plants averaged 0.85 g of root DW per gram of shoot DW (including all aboveground parts plus the crown), whereas in plants fertilized with 50 and 100 kg·ha" N, the ratio averaged 0.61 and 0.48 g.g", respectively. By contrast, the second year, the ratio was similar among treatments, averaging 0.34 to 0.39 g-g" at fruit harvest. Root-to-shoot ratios often decline with N fertilizer application and plant Accumulation and partitioning of plant age (Gregory, 2006); however, excavation of biomass. Plants fertilized with 50 kg·ha" N the root system also becomes more difficult as were the largest among the three N treatments the plants grow and mature. Such difficulty and, by the end of the first growing season, likely reduced the percentage of roots recovproduced 0.36 kg of total DW and weighed ered in the present study and perhaps partlY 22% more than plants fertilized with 100 accounted for the lower root-to-shoot ratios kg·ha-' N and 32% more than unfertilized and the lack of treatment differences observed plants. By fruit harvest the next season the during the second year after planting. difference was even greater as plants' ferThree major losses of DW occurred durtilized with 50 kg·ha" N averaged 0.70 kg ing the first 2 years after planting. The first of total DW per plant, whereas those with was at leaf abscission in early to mid-Nov. 100 kg·ha" N or no fertilizer averaged only 2002. Leaf abscission resulted in a net loss of 0.~9 and 0.37. k~plant, respectively. Clearly, 43 g of DW or 16% of the total biomass in SOli N was lImIted at the site, but adding unfertilized plants, 92 g or 25% of the total ~ 00 kg'h~" N was excessive for this plant- biomass in plants fertilized with 50 kg·ha" N, mg denSIty and resulted in reduced plant and 86 g or 29% of the total biomass in pla.nts gro\~h [see Baiiados et al. (2012) for further fertilized with 100 kg·ha-' N. At this pomt, detaIls and additional harvest dates]. Re- fertilization with more N resulted in a greater duced gro\\1h with excessive N fertilization allocation of biomass to leaves and therefore 68Ilg·g" with no N to 30 Ilg·g" with 50 kg·ha-' Nand 28 IlgK' with 100 kg·ha-' N. LeafB was not deficient (less than 20 Ilg·g") in either year, whether plants were fertilized with N or not, but levels were slightly less than optimum (i.e., 31 to 80 IlgK'; Hart et aI., 2006) the second year after planting when N fertilizer was added. Leaf concentrations of the remaining nutrients, including K, Fe, Cu, and In, were unaffected by N fertilizer application and, with the exception of Fe (Fig. IH), remained more or less the same during the first 2 years after planting. LeafK concentration averaged 0.78% over each treatment and year, and leaf Fe, Cu, and In concentrations averaged 333, 5, and 9 Ilg'g-', respectively. Both Cu and In were within a range considered adequate for these nutrients (5 to 15Ilg'g" for Cu and 8 to 30 for IlgK' for In; Hart et aI., 2006), whereas K and Fe were higher than adequate (i.e., 0.41% to 0.70% for K and 61 to 200 Ilg·g" for Fe; Hart et aI., 2006). Although others have reported an increase in leaf K concentration with N fertilization in highbush blueberry (Bishop et aI., 1971; Cwnmiogs, 1978; Cwnmiogs etaI., 1971; Townsend, 1973) and an increase in leaf K as plants aged (Cummings, 1978), fluctuations in leafK are often associated with fruiting; these plants produced no fruit in Year I and only a small crop in Year 2. Table I. Effects of nitrogen (N) fertilizer rate on flower and fruit dry weight and macro- and m' t' . . dyear after planting.' Icronu nent content m 'Bluecrop' blueberry dunng the secon N rate (kg-ha") Flowers 0 50 100 Significance' Dry \\1 (g/plant) 4.1 4.2 3.1 1'>S N Macronutrient content (g/plant) Phosphorus Potassium Calcium Magnesium 0.13 0.21 0.16 0.02 0.03 0.02 0.06 0.08 0.06 0.02 0.02 1'>S 1'>S NS NS om Micronutrient content (mg/plant) _ Boron Copper Manganese _~ Sifu u r Iron om 0.01 0.02 0.01 0.8 1.4 1.0 0.4 0.2 NS NS NS 0.01 O.QI OJ 0.2 0.1 0.1 1.1 3.9 2.7 NS NS NS OJ OJ 0.2 ss Fruit' 116 0.88 0.10 0 0.80 0.05 0.04 0.7 0.08 3.8 4.2 0.9 2.34 171 0.3 0.15 50 1.19 0.07 0.Q7 1.0 0.12 5.6 6.1 1.4 0.88 55 0.05 0.4 100 0.37 0.02 0.02 OJ 0.04 1.7 \.9 0.4 Significance' ~ 0.1 'Flowers were collected on 22. Apr. 2003, and fruit were harvested on 25 July 2003. 'Results from one-way analYSIS ofvanance: NS and § = nonsignificant and P < 0.10 res t' I . ' content except N . the average of2 years of data 'collect pec dIve 'All fruit nutnent I was ca Icu Iate d usmg f iY ' , . d rom 200 kg·ha" N (unpublished results). The plants were growing at the same site and sampl de th mature Bluecrop' plants fertilized each yearWlth 0, lQO,an e e same years as those used m . the present study. 920 HORTSCIENCE VOL. 47(7) JULY 2012 a relatively greater loss of total biomass at leaf abscission. The next loss of OW occurred during winter pruning in which 28 g of wood was removed from unfertilized plants, but only 9 and 4 g of wood were removed from plants fertilized with 50 and 100 kg·ha- 1 N, respectively. Unfertilized plants required more pruning than fertilized plants because this treatment had a lot of weak and twiggy wood. Twiggy wood typically has thinner and shorter new growth and produces small berries and therefore is often removed during pruning (Strik et aI., 1993). The final loss of OW occurred at harvest when an average of 55 to 171 g of fruit per plant was picked (Table 1). Although production was variable, plants fertilized with 50 kg·ha- 1 N tended to produce more fruit than those fertilized with 100 kg·ha- I N. Plants also lost a small amount of DW (less than 3 to 4 g/plant) at fruit set as the flower petals senesced (Table I). Accllmlilation and partitioning ofnlltrit'lIIs. In most cases, nutrient content in each plant part followed the same pattern of accumulation as ow (Figs. 3 to 6). Thus. many differences among N treatments were similar. For example, nutrient content in new shoots ...• 0 kg'ha-' N --0- 50 kg'ha-' N -...- 100 kgha" N 'E rn a. :§ z A Year 1 3 N' T" NxT" iii 2 ""-0 B 4 Year 2 Year 1 ~ a T- o "a a NxT" // a T" NxT" a p/ 2 .. /a / I. . b 0 .s:: en ~ iii 2 ""-0 0 .s:: en 1.2 1.0 0.8 C / ~ / / a r' / / / 0.6 • 0 /. 0.4 I b /. 0.2 a Year 1 P NNS b b • I a 1.2 1° 1.0 /b 06 • 06 •c I I 'b O> 0.3 :e-.!2l ~ iii 2 0.2 ""-0 0.1 0 .s:: F E 0.4 r:rT" NxT ~ TNxT" ~ TNxT" NS :e0> E. 60 50 Q) 40 'lii 30 ""-0 20 .s:: 10 u. 2 0 en (f) ;:r 5? 00 10 a '" iO () 0.4 -0 or 0.2 c::> /. 0 ~ TNxr ~ ta • b • I 1 Year 1 r:rT" ... I b ~ - 0.2 • NxT NS • fi .' / /.. a I ! 0 /, 1 /. ;, N' TNxT- 0.0 /b ::r 6 ~ 6 10 00 a /56 b I / /, or (J) NXT-g ___ ~ ? /, i€: '0 6 b Year 2 T" a a (f) I N' N" T" NxT" 00 I !. ~ 10 0.4 I Year 1 Year 2 ;:r 0.6 1a H G (f) a Year 2 a 0.0 rn a Year 1 Year 2 Year 1 en 'E ~ 0.0 0.0 'Ern ::!. ~ 0 I I 1 . b • /, h /. ~ T" NxT" NxT- 0.05 ~ a 0.00 Year 2 T" 1 1 a 1 1 1 / NxT" • / /. 0.15 0.10 / o a }) Year 1 p / 020 / a / 0 1.4 a N' T" NxT' .-0 a en Year 2 ~ / I / •b CIl 4 ,t! '3 <fl. 2 /, / a "C /if .5 0 0 a 'E rn ~ E. Year 1 120 N' T" NxT- c ~ 'lii 2 ""-0 0 .s:: 90 I /. ,$ /. .b July I I I I. b • Oct Jan_ Apr. Year 2 T" N" T" NxT' / I ..- ,0 a Year 1 ~ /a I a a , 30 0 Apr. N" T" NxTI 60 en I Year 2 (J) J 0 150 p-:/ • /. b • /, Apr. I la f I. / July a /, Oct July Jan. ~ 2.0 "" Apr. 1.5 1.0 b ::r 2.5 }) 0.5 1D ~ N ::> '3 '"-0or ~ 00 July 2002-2003 . 2002-2003 . Fig. 3. New shoot and leaf A) nitrogen (N), (B) phosphorus.(Pl· (C) po~s~;~:;~ K D I'urn (Ca) (E) magnesium (Mg). (F) sulfur (S). (G) Iron (Fe). (HI first (2(io2) and the se.cond (2003)year ?fter planting: )'JuJn~~~ The.pl;~~ ~oronf,(8~, .(1) mangane~~ (M?), and~ z;n~o(~;.~~~~~~':s!~u::~Ptwo-way~alysis o~\':::~~~:e~~;a:~~e~~~:~ i~~~~tl~~:~~::~~h :~o~mon J) n:re. er.tfilhzed each.sp,:,ng with ~ ~ ~~ and 001 respectively. Each symbo! represen~tt. ~ range test) Shoot and leaf copper (Cu) content increased to an nSlgnl cant and Significant at - . . - , t th 50!. )evel (Duncan s new mu Ip e . letter on a given date are not significantly ddfen:nt ~ e b~ was unaffected either year by Nor N x T. 2 average of 0.6 g/plant in Year I and 0.7 glplant III ear 921 0 HORTSCIENCE VOL. 47(7) JULY 2012 lEaLI content when no fertilizer was applied (Figs. 4C, 4H, and 41). In addition, the content of several nutrients in the crown differed among N treatments, in which N, P, S, and Zn content was higher at times with N fertilizer (Figs. 5A, 5B, 5F, and 5J), whereas K and Ca content was sometimes lower (Fig. 5C-O). Basically, differences in root N, P, K, Mg, S, Cu, and Mn content were similar to differences in root 3J), largely as a result of a higher concentration of each in unfertilized plants (P :::; 0.05). Another exception was that wood Nand S content was higher in fertilized plants much earlier than OW, and this occurred whether 50 or 100 kg·ha- 1 N was applied (Figs. 4A and 4F). Wood K, Cu, and Mn content also differed among treatments before harvest and, in one instance, wood had a higher Cu and leaves was generally higher each year in fertilized plants than in unfertilized plants (Fig. 3). Likewise, like with OW, the content of many nutrients in wood and roots was often highest in fertilized plants (Figs. 4 and 6). However, there were a number of exceptions. New shoot and leafMg, Fe, and Zn contents, for example, were similar among treatments the first year after planting (Figs. 3E, 3G, and ... o kgha" N -D- 50 kg·ha" N - y - 100 kg·ha" N c !!l "'- ~ 1.2 0.9 Pruned A N" T" NxT- a 0 0 ~ 0.6 0.3 b 0.0 0.30 C 0,25 '" ~ Ci ~ -0 0 0 ~ 0.20 .. b • Pruned 0.12 t a ./ .. Z -0 B N' T" NxT' t ./ ./ • •c ... -D b C . 9 Pruned N- t r' NxT" / / ab , a ... a / • l· ~ .. '\ .. I.e '\ \-, l D N' T" NxT NS .. b { t.. ::;: '" 0.03 -0 0 0 ~ 0.02 € 0.04 iil i': -0 0.02 0.25 0.20 ~ 8. (') /, 0.15 '" /- <0 h 0.10 0.05 0,00 0,04 0.06 Pruned 0.05 ~ 0 0.00 0.10 0.06 ~ 0.08 ~ b a 0,15 C !!l 0.05 0.10 ~ i': 0.00 Pruned E F t NNS • T" NxT- ........• • • 0.01 Pruned 0.12 a - ,,9 _~// b a • ~ g a. 0.095!! / / /0 .....•.......... 0,00 0.15 a a b ,9 t N" TNxT" co -0- .. a • b b 0.06 iil •c b i': 0.03 0.00 2.5 C '" Ci 0, £ 2.0 G H NNS NNS T" NxT NS Pruned t }J-., 1.5 CD -0 0 0 ~ 1.0 I. & .. • TNxT- Pruned t 12 a • a /0 /:. .e_ ........ 0.5 _-0 ,/ .. b 0.0 b ~ 10 0 0 8 C 6 4 2 a. (') 3' co -0iil :> .0 0 a C 80 !!l a. 0, 60 £ o ,, NTNxT' Pruned ta 40 / 20 // A • ....... ab..... I / / / , J Pruned t 0 I b I l- • ··b I • 2.5 ~ 2,0 "'- 1.5 3' 0 1.0 b··. 0.5 July Oct Jan. Apr. July 0 N :> ce. -0 iii' E: AP;r.~~J-;:-u:;:-Iy-:-~o~c'::t~,~-:-JaJ..n-,~---,-~~-,-...J 0.0 Apr. 2002-2003 July 2002-2003 Fig, 4. Woody stem (Al nitrogen (N), (B) phosphorus (p), (C) potassium (K) (D) I' (Cu,l., (I) manganese (Mn). and (J) zinc (Zn) content in 'B\uecrop' bluebe~ d:n~l~~ (Ca), (El magnesium (Mg), (F) sulfur (S), (G) boron (B), (D) copper fertlhzed each spnng WIth 0,50, or 100 kg·ha- ' N. Results from two-wa I ': first.(2002) and the second (2003) year after planting. Theplant5 \\ere " f icant an d' Y ana YS1S of vanance [N rate (N) x time (1)] are inset in each graph: NS, • ' ., '" nonslgm slgm'ficant at P --< 00. ) an dO .01, respectively. Each symbol Ietter on a given . date are not slgm ' .ficantIy d'Iflierent at the 5% level (Duncan's . represents the meano fthree replicates; means without letters or with a coromon new' but was unaffected by Nor N x T. multiple range test). Wood iron (Fe) content ranged from 4 to 17 mglplant 922 HORTSCIENCE VOL. 47(7) JULY 20P~ ..• 0 kg·ha-' N --0- 50 kgha-' N 1.:-_____---:-____----'Y-_~100 kg·ha-' N 0.7 ~--------------- A a :g 0.6 N' 0. '" 0.5 NxT' z 0.4 ~ 0.3 E? c ~ B a P' .. a / .... 0.2 .............• a 7 • b b 0.14 0.12 :e-'" 0.10 ~ '" ~ 0.04 ~ 0.00 D C NNS T" NxT' N' P' NxT" a • 0 / r-.. 0.06 0.04 0.02 n a 0.05 '\. ... . ... ~ 6 0.08 ~ 0.06 ~ :> 0.02 00 :g n 0.08 ~ 0.062' b 0.1 0.10 ~ 0.04 n OJ 0.03 iO -0 0.02 ~ 0.01 ~~~~-'---'~---'-~-'---~-'--.....J 0.00 _ 0.05 c: ~ 0.04 E? .. .. NNS T- NxT NS ./ / ... ~ 0.02 e u 0.01 <II ~ 1.5 1.2 §. 0.9 C e u NxT" / "a .... • H G P' :> 0.06~ 0.04 '"~ NS -.""'/ e ~ 3 (') c 2 ]' -0 0.6 ii> :> • .... 0--"- 0_3 .a 4 n NNS TNS NxT NNS NxT (') 0.08 ~ b 0.02 b '--"-~J-~-L~~'---'-~--'--~-'------J 0.00 ID ~ 0.10 a T- p ~ 0.03 :g F N' E .e 0.0 J :g 30 <II E- 25 P' §. 20 NxT O) c :::; NNS NS '/ V 15 'P • ~ 7 6 /,a 7 a .. c U a 0 (') N' TNxT' 10 5 4 • b 3 2 ~:> N :> '3 ~OJ ~ b 5 0 July Oct Jan. Apr. 2002-2003 July Apr_ July Oct Apr. Jan. July 2002·2003 r 19.~. Crown (A) nitrogen ~, (8) phosphorus (P), (C) potassium (K), (D) calcium (Ca), (E) magnesium (Mg), (f) sulfur (S), (~) boron (B). (II) copper (Cu)' (ll angan~se (Mn), and (J) Zinc (Zn) content in 'Bluecrop' blueberry during the first (2002) and the second (2003) year after planll1lg. The plant~ were fertilized each ~~g WIth 0,50, or 100 ~.ha-l N. Results from two-way analysis ofvariance ~ rate (N) x time (1)] are inset in.each graph: NS, ., •• ~ ~ignificant and significant at di - 0.05 and 0.01, respectively. Each symbol represents the mean of three replicates; means "lthout letters or Wlth a common letter on a given date are not significantly fferent at IDe 5% level (Duncan's new multiple range test). Crown iron (Fe) content ranged from 3 to 29 mg/plant but was unaffected by Nor Nx T. DW, although sometimes the differences occurred in the 50 and 100 kg·ha- 1 N treatments anfted, unlike root DW usually occurred more o ntha . 6F 6H, nJust at fruit 'harvest (Figs. 6A-C, 6E, , and 61) The conten~ of many nutrients in the Woody . (29 0 stems lIlcreased over fall and winter . ct. 2002 to 7 Feb 2003' Fig 4A-J) The IDcreasf .,.' tre trn eoN, P, K, and Mg in almost all a ents from fall to late winter may have been a result of remobilization of nutrients from senescing leaves or additional plant uptake during this time period. Copper content likely increased as a result of a copper sulfate fungicide (see subsequently). The content of Ca in the woody stems increased over fall/winter; because this nutrient is not mobile, it is unlikely this increase was from remobiIization before leaf abscission. In the crown tissues, only N, P, and K content in the unfertilized plants increased over winter, perhaps a result of added nutrient uptake during this period. In the roots, N content increased in the unfertilized plants and P in those fertilized with 100 kg·ha- ' N, perhaps a result of the lower soil pH in this treatment (Baiiados et al., 2012). which would have increased availability of soil P. Losses in nutrient content over this time period may have been caused by a reduction in efficiency 923 HORTSCIENCE VOL. 47(7) JULY 2012 • 0 kg'ha- 1 N -{)- 50 kg'ha- 1 N -...- 100 kg'ha- 1 N A 1:: 3 m a. :§ z a N" P" / / ~< 1) 0.15 b / I. • • / b b /. 0 0.5 1:: m a. :§ ><: (5 a rx: 0.4 ;0 / / / / ....... ~--- 0.25 0.20 ~ / / a (5 / NxT" / 2 0 TO> / a NxT"" rx: o ~--------------I B a P N' <0 -0 0.10 ~ c 0.05 0.00 a C D N' o / N" T"' NxT' I T" / / / .• -- .>......, • 1 NS ." I b / ..... ......... . . . /. 0.1 1.2 ;0 0 1.0 .,(') 0.8 0---"9-.. b / I' 0.2 NxT / r, 0.3 1.4 ..... 0-- •. ~ 0.6 <0 -0 0.4 c" or 0.2 0.0 0.5 'E -[ 0.4 :§ '" :2 g 0.0 E N" F NO. a T" NxT' / fa / NxT" a / 0.3 0.8 a T" ? a / / 0.2 / rx: .... b 0.1 c '" a. en .s ttl (5 4 :::J c • b 0.2 0.0 H NNS NNS TO> 3 C/l or / /, t ~ 2- <0 0.4 -0 / b G ;0 0 0.6 o NS NxT NxP' J. 2 /, a a p T" I I • rx: 0 2- 6 '3 <0 4 • • b b ::0 8 () c I i· 10 -0 or :::J 2 c 0 150 C a. '" en 120 .sc 90 (5 60 :2 J N" T"" NxT" I 0 rx: lab I 30 • b 0 Apr. I a A, July Oct , , 2002-2003 a 0 15 b 6 3 Apr. July 2N :::J 9 I I ::0 a 12 I .. Jan. NNS T" NxT NS Apr. July Oct Jan. Apr. '3 <0 -0 or Eo 0 July 2002-2003 Fig. 6. Root (A) nitrogen (N), ~B) phosphorus (P): (C) potas,sium (K), (D) ~alcium (Ca), (E) magnesium (Mg), (F) sulfur (S), (G) boron (B), (H) copper (CW~~J manganese (Mn), and (J) zmc (Zn) content m Bluecrop blueberry dunng the first (2002) and the second (2003) year after planting. The plants were .fe~ t I each-spring with 0,50, or 100 kg·ha- N. Results from two-way analysis of variance [N rate (N) x time (1)] are inset in each graph: NS, *, ** = and significant at P ~ 0.05 and 0.0 I, respectIvely. Each symbol represents the mean of three replicates; means without letters or with a common Ie e a give~ date are not significantly different at the 5% level (Duncan's new multiple range test). nonslgn~fi~~n of harvesting the entire root system as plants 1 aged. Like DW, plants fertilized with 50 kg·haN tended to have a higher fruit N content (Table I). As mentioned earlier, other fruit nutrients were not measured, but measurements done on fruit from mature 'B1uecrop' plants gro\\ing at the same site an~ f~ilized with 0, I 100, and 200 ka·haN indIcated :;, . that . N treatment had no effect on frUIt nutnent concentrations beyond fruit N [which increased with N fertilization (1.02% to 1.28% N; Baiiados, 2006), as previously reported by others (Ballinger and Kushman, 1966; Bishop et aI., 1971)]. In this case, fruit averaged O.W/o P, 0.69% K, 0.04% Ca and Mg, 0.07% S, 33 I1g-g-1 Fe, 8 I1g'g- 1 B,2 Jlg.g-I Cu, 36 I1g-g-1 1 Mn, and 6I1g'g- Zn (Strik et aI., unpublished data). Assuming nutrient concentrations were similar in the present study, the fruit would have contained 0.9 to 2.3 g/plant 0 fN·04to , p' ell, 1.2 g/plant of K; less than 0.2 g/plant of : and Mg, and S; 2 to 6 mg/plant of Fe and ~i I) less than 2 mg/plant ofB, Cu, and Zn (Ta ~ ; With the exception of Cu, total nu~e~ uptake by the plants increased from n:it budbreak to leaf abscission (Year I~ O~any harvest (Year 2) and was higher. m Fig. cases when plants were fertilized With N ~ IS 7). With N fertilizer, uptake of some nutnen 924 HORTSCIENCE VOL. 47(7) JULY 2012 ...• 0 kgha" N -()- 50 kgha" N 'E 10 .e- 8 z 6 .!!1 S C r:-__________-_T_- _100 kgha" N r---------------a A \) B N" ]j 0 I- .. 2 .....•. b '" i'i :9 ~ C / •c b o p P' N' r' NxP' NxP' N' I p/ / b :::< C '" i'i ~ I- ~ b 0.2 E ;a / .c 1.0 <0 0.5 .5: I r" 'i2: ., I .. I .... I 1.0 .. • b 0 __ I 9 I I / .. t 9 a I) a U 0.5 ~ / 400 0, I 300 / N" ? / r" / NxP b I I • I Jan. 2002-2003 Apr. July Apr. July o ab / I Oct. 10 3" S a J I July ";aOJ c: a ~ 100 15 <0 b i'i [ (") b 3 f,j 200 c} 20 / / • . I :::< ~ <0 b I t· 6 ~ "OJ b I " Pruned '\ a NxP' I t / : ">;;:;a I 1.5 P' I lab" -4 N" I a Q. NxP' 9 Pruned c ., (") F a N" 12 S OJ 1.5 b c -[ 0 b / ..... -I [ 0.0 0.6 0.4 2.0 I I I 0.8 u-OJ .5: -0 I OJ <0 I 0 '" ~ -0 0.0 '" ~ I- i'i ;a a 2 1.0 0.4 0.2 i'i 'E OJ b / / C 3 0 [ -0 a.···· c 0 'E .. a 4 ~ b 0.6 I Pruned NxP' a <II i'i I r" NxP' -I I N" r" Oct Jan. Apr. UOJ .5: -I 20 [ 15 ";aOJ 10 ]" N •b S " UOJ .5: July 2002-2003 ,g'(i) Total plant (A) nitrogen (N), (B) phosphorus (P), (C) potassium (K), (D) calcium (Ca), (E) magnesium (Mg), (F) sulfur (S), (G) boron (B), (II) copper (Cu). F' fe m,anganese (M~), and (J) zinc (Zn) content in 'Bluecrop' blueberry dunng th~ first (2.0.02) and the second(2003) year .after planting. The plants were n r!tl~ze? each spnn: with .0, 5.0, or 1.0.0 kg.ha- 1 N, Resul~s from two-way analYSIS of vanance [N rate (I") x. tIme (nJ are tnset III each graph: M. *, .. ~ g I onSlgnlficant and slgmficant at P:=; .0..05 and .0,.01, respectIVely. Each symbol represents the mean of three rephcates; means WIthout letters or with a common cttcr on a given date are not significantly different at the 5% level (Duncan's new multiple range test). Such . . y as Ca" S an d B was general1y higher III (:ar I at the beginning of the growing season OcPnl to July) than later in the season (July to tober), whereas uptake of other nutrients M M . suchasK,g, th n, and Zn was ' higher later III e season. However when no N fertilizer ' was limited later in wasaPP rled, plant growth th nu~ .seaso~ (Fig. 2), and uptake of many nents, Including N P K, Ca S and B Was " " , Unl~educed, and only Cu uptake was higher. e other nutrients, Cu uptake also increased over winter between the first 2 years after planting, whether N was applied or not (Figs. 4 and 7). The plants were sprayed with copper sulfate fungicide (Bordeaux) after pruning, a common practice in blueberry to control bacterial canker (Strik et aI., 1993). Loss and net gain of nutrients. Loss and net gain of nutrients in plants fertilized with 50 kg·ha- ' N are summarized in Table 2. Total nutrient losses, including leaf abscission, pruning, and fruit harvest, averaged 4.85, 0.28,2.38,0.82,0.25, and 0.54 glplantofN. P, K Ca, Mg, and S, respectively, and 5U, 5.20, 0.57, 42.4, and \.71 mglplant of Fe, B, Cu. Mn, and Zn, respectively. At a density of2691 plantslha, each loss was equivalent to 13.1 kg·ha- ' N, 0.8 kg·ha- ' P, 6.4 kg,ha- 1 K, 2.2 kg·ha- 1 Ca, 0.7 kg·ha- 1 Mg, 1.5 kg'ha~l S, 138 g·ha~l Fe, 14 g·ha- 1 B. \.5 g·ha- ' Cll, 114 g.ha- 1 Mn, and 4.6 g·ha- ' Zn. More than half the N, P, K, Cu, and Zn was removed during fruit harvest, whereas most Ca, Mg, S, Fe, M;. 925 HORTSClENCE VOL. 47(7) ~ JULY 2012 Table 2. Loss and net gain of macro- and micronutrients in 'Bluecrop' blueberry during the first 2 years after planting.' Macronutrients (g/plant) Potassium Calcium Magnesium Sulfur IronY Micronutrients (mg/plant) Boron Copper Manganese Zinc Loss 0.09 38.2 3.55 0.39 24.1 0.17 0.69 OJI 1.09 Leaf abscission' 0.10 2.18 7.3 0.25 0.03 12.2 0.01 0.06 0.40 0.10 Pruning 0.04 0.33 lAO 5.6 0040 0.12 6.1 0.07 0.07 1.00 Fruit harvest 1.19 2.34 0.15 Gain 2.38 39.2 1.36 0.50 145 0046 ll.8 0.24 0.92 Bud break to leaf abscission (Year I r 0.56 2.55 7.36 6.64 1.16 19.3 285 0.69 lAO 10.5 Bud break to harvest (Year 2)" 0.33 1.69 5.54 'Plants were fertilized each spring with 50 kg·ha- ' nitrogen. 'Net iron gain does not include iron in the roots. 'Estimated from leaves collected on 29 Oct. 2002. ·Calculated as the difference in wood sampled just before pruning (7 Feb. 2003) and wood sampled at budbreak (22 Apr. 2003). 'Calculated as the difference in nutrients of whole plants (minus any leaf nutrients) sampled just before leaf abscission (29 Oct. 2002) and whole plants sampled at budbreak (29 Apr. 2002). "Calculated as the difference in nutrients of whole plants (minus any fruit nutrients) sampled at harvest (25 July 2003) and whole plants (minus any flower nutrients) sampled at budbreak (22 Apr. 2003). Nitrogen Phosphorus om W and B was lost during leaf abscission. Although the nutrients in senesced leaves may be lost from plants in the short term, if leaves drop into the row, the nutrients would likely be available to plants again once leaves decompose (Strik et a!., 2006). The relative losses of nutrients were similar in plants fertilized with 0 and 100 kg·ha- I N, although unfertilized plants (which allocated 32% total biomass to fruit) tended to lose proportionally more nutrients during harvest, whereas plants fertilized with 100 kg·ha- I (which allocated only 13% total biomass to fruit) lost most nutrients during leaf abscission (data not shown). Net nutrient gains averaged 8.09, 0.57, 2.25,2.32, LIS, and 1.66 glplant ofN, P, K, Ca. Mg, and S, respectively, and 58.5, 9.74, 8.00, 431, and 22.3 mg/plant of Fe (not including roots), B, Cu, Mn, and Zn, respectively (Table 2). When combined with the losses, plants fertilized with 50 kg·ha- I N required the equivalent of 34.8 kg·ha- I N, 2.3 kg·ha- I P, 12.5 kg·ha- I K, 8.4 kg·ha- ' Ca, 3.8 kg·ha- I Mg, 5.9 kg·ha- I S, 295 g·ha- I Fe, 40 g·ha- I B, 23 g·ha- I Cu, 1273 g·ha- I Mn, and 65 g·ha- I Zn over 2 years. Thus, the relative amount of N, P, and K fertilizer used during establishment, including the N added during sawdust incorporation and in the nursery potting media at planting, was 21%, 3%, and 9%, respectively. Because plants were smaller, fewer nutrients were required when 0 or 100 kg·ha- I N was applied and therefore percent fertilizer used in these treatments was even less. This study has generated the first comprehensive nutrient accumulation data set from planting to fruit harvest for establishing highbush blueberry. The information was used to determine nutrient requirements of the young plants, a useful tool for developing efficient fertilizer management practices during the critical stage of field establishment. Plant uptake of most nutrients increased throughout the growing season, indicating that it is best to apply the majority of fertilizers in the spring; however, K. Mg, Mn, and Zn uptake was higher later in the season and therefore it may be better to apply these nutrients, ifneeded., in early or midsummer. Next, we plan to evaluate nutrient requirements in relation to N supply in mature plants. Literature Cited Abbott, J.D. and R.E. Gough. 1987. Seasonal development of highbush blueberry under sawdust mulch. J. Amer. Soc. Hort. Sci. 112: 60--62. Ballinger, W.E. and L.J. Kushman. 1966. Factors affecting the mineral-element content ofleaves and fruit of Wolcott blueberries. Proc. Amer. Soc. Hort. Sci. 88:325-330. Baiiados, M.P. 2006. Dry weight and 15N-nitrogen and partitioning, growth, and development of young and mature blueberry plants. PhD diss., Ore. State. Univ., Corvallis, OR. Baiiados, M.P., B.C. Strik, D.R. Bryla, and T.L. Ri~etti. 2012. Response of highbush blueberry to rutrogen fertilizer during field establishmentI. Accumulation and allocation of fertilizer nitrogen and biomass. HortScience 47:648655. Bishop, R.F., L.R. Townsend, and D.L. Craig 1971. ~ffect of source and rate of N and M~ on nutnent levels in highbush blueberry leaves and fruit. HortScience 6:37-38. Cummi~~s, G., C. Bickford, and L. Nelson. 1971. FertIlIzer and lime rates influence highbush blueberry groMh and foliar elemental content dunng establishment. J. Amer. Soc. Hort Sc' 96:184-186. . 1. Cummin?s, G.A. 1978. Plant and soil effects of fertl!lzer and lime applied to highbush bluebemes. J. Amer. Soc. Hort. Sci. 103:302-305 Gre?ory, P.. 2006. Plant roots. GrOMh, activity and mteracllon With SOIl. Blackwell Pub!" h' Ames, IA. 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