The effect of soil pH on plant growth, leaf chlorophyll

fluorescence and mineral element content of two
blueberries
Y. Jiang, Y. Li, Q. Zeng, J. Wei and H. Yua
Institute of Botany, Jiangsu Province and Chinese Academy of Sciences, Nanjing, China.

Abstract
Soil pH is a key factor affecting the growth of blueberry, and most soil in China is
difficult to grow blueberry because of the high soil pH. Therefore, the study on the
reaction mechanism of blueberry to different soil pH is beneficial to breeding new
cultivars to adapt to different soils with excessive pH. Research was conducted to
study the effect of different soil pH (4.5, 5.3, and 6.0) on plant growth, chlorophyll
fluorescence, and leaf mineral element content in two bluebrerries, Vaccinium
virgatum ‘Climax’ and V. corymbosum A119, an excellent seedling progeny from the
southern highbush blueberry cultivar ‘Southmoon’. Plant height, stem-base diameter,
branching number, leaf dry weight, stem dry weight, root dry weight, chlorophyll
content index, PSII maximum photochemical efficiency and effective light quantum
yield showed decreasing trends with increased soil pH in the two blueberries.
However, less reduction was found in A119 than ‘Climax’. Leaf Fe and Mn contents of
‘Climax’ and A119 decreased as the soil pH increased, whereas K, Ca and Mg contents
increased. P contents decreased in ‘Climax’ and remained fairly constant in A119 as
the soil pH levels increased. Cu and Zn contents were unaffected irrespective of pH in
the two blueberries. In total, A119 had greater high pH tolerance than ‘Climax’, and
the disorder of leaf mineral element content could be a key factor to inhibit the
growth of blueberry when the soil pH was increased.

Keywords: soil pH, blueberry, growth, leaf nutrient content, chlorophyll fluorescence

INTRODUCTION
Blueberry (Vaccinium spp.) is an acid-loving plant that grows in a soil pH of about 4.0-
5.5 (Harmer, 1944). Blueberry growth is inhibited and leaves can become chlorotic due to
iron deficiency under high soil pH (Holmes, 1960), which results in the decline of fruit yeilds
(He et al., 2009). The soil pH of most blueberry fields is not suitable for its proper growth, so
this often requires artificial soil acidification to grow blueberries (Haynes and Swift, 1985).
However, artificial acidification of soil is too expensive, and the soil pH will still rise after a
period of time, resulting in poor growth and low production of blueberry (Finn et al., 1993).
Planting high pH resistant blueberry varieties is a key way to solve the problem.
Different blueberry genotypes have different adaptation to high soil pH. Finn et al. (1993)
found variation was significant among thirty-three seedling progenies from crosses among
highbush blueberry (V. corymbosum), lowbush blueberry (V. angustifolium), and V.
corymbosum × V. angustifolium hybrid-derivative parents. ‘Northblue’, ‘Northsky’, and
‘Northcountry’, and V. angustifolium were not a general source of tolerance to higher pH, but
some specific populations derived from V. angustifolium were tolerant of high soil pH. Tsuda
et al. (2014) studied the response of four hybrid clones between V. corymbosum ‘Spartan’
and V. bracteatum to high pH, and found the rooting rates of one clone, JM4, (91%) at pH 8.0
was significantly higher than ‘Spartan’ (0%), and indicated JM4 had higher pH tolerance.
In the south of China, the rabbiteye blueberry (V. virgatum) and the southern highbush
blueberry are the most common types of blueberry (Yu et al., 2012). However, the fruit
ripening period of rabbiteye blueberry is during the rainy season in this area (He et al.,
a
E-mail: njyuhong@vip.sina.com

  Acta Hortic. 1180. ISHS 2017. DOI 10.17660/ActaHortic.2017.1180.36 269

Proc. XI International Vaccinium Symposium
  Ed.: J.W. Olmstead
2009), which resulted in the smaller fruit and dehiscence, and a decrease of sugar content
and yield. The fruit maturation period of southern highbush blueberry is about one month
earlier than that of rabbiteye blueberry, which can avoid the rainy season. This results in
larger, better tasting fruit suitable for fresh consumption. However, the weaker adaptability
of southern highbush blueberry compared with rabbiteye blueberry resulted in poor growth
and low yield and quality, and limited its use in this area (He et al., 2009).
Through breeding, some excellent selections with high yield, good quality and high
adaptability were obtained from ‘Southmoon’ southern highbush blueberry at the Institute
of Botany, Jiangsu Province and Chinese Academy of Sciences (Zhang et al., 2009). Here, we
used one selection, A119, and one cultivar, ‘Climax’, a rabbiteye blueberry widely used in this
region, to study the effect of different soil pH on plant growth, leaf nutrient content,
chlorophyll content, and chlorophyll fluorescence characteristics to provide experimental
basis for further selection and breeding of blueberry cultivars with high adaptability to high
soil pH.

MATERIALS AND METHODS

Plant materials
Two blueberries including A119 southern highbush blueberry and ‘Climax’ rabbiteye
blueberry were used in this study. A119 is a seedling selection from ‘Southmoon’ with high
yield, good quality and high adaptability, which was selected by the Institute of Botany,
Jiangsu Province & Chinese Academy of Sciences, China. ‘Climax’ was a frequently used
cultivar in Jiangsu Province, China and was used as control in this study.

Experimental methods
The experiment was carried out from April 13, 2013 to September 15, 2013 in a
plastic greenhouse under natural conditions. Daily maximum and minimum greenhouse
temperatures were presented in Figure 1. The maximum photosynthetically active radiation
about ranged from 600 to 2000 µmol m-2 s-1. Uniform, one-year-old plants of each blueberry
(about 20 cm high) were planted in plastic pots (diameter 20 cm, depth 20 cm) filled with 5
L mixed matrix of 2 acidic red soil:2 peat:1 perlite (by volume). Before planting, NH4NO3 8.10
g, Ca(H2PO4)2·H2O 5.06 g, and K2SO4 5.25 g as a base dressing was added to each pot and
mixed. The soil was treated with different rates of CaCO3 to give final pH values of 4.5, 5.3, or
6.0. One plant was planted in a pot as one replication, and four replications were used for
each pH treatment. After planting, they were irrigated with tap water. Afterwards, plants
were watered to maintain 65-75% maximum field capacity.

Figure 1. Daily maximum and minimum temperatures in the greenroom.

After growth in different soil pH for approximately 5 months, plant height, base-stem
diameter, and shoot number of each plant were measured. Then the plant was taken out
from pot, and the harvested plant materials were washed 3 times using tap water, followed

270
by 3 times using double-distilled water. Plant materials were then divided into roots, stems,
and leaves, and oven-dried at 80°C for 48 h; dried weights of different plant parts were
measured and recorded.
Before harvest, the chlorophyll content of newly formed leaves was analyzed with a
chlorophyll meter (CCM-200, OPTI-SCIENCES, USA), and the chlorophyll content was
recorded as a chlorophyll content index. Chlorophyll fluorescence (PSII maximum
photochemical efficiency and effective light quantum yield) of newly formed leaves were
also determined with PAM chlorophyll fluorimeter (OS1p, OPTI-SCIENCES, USA). Five leaves
of each plant were used for chlorophyll content and chlorophyll fluorescence determination.
Dried leaves were ground to a fine powder, and 0.2 g was digested in a mixture of
HNO3:HClO4 (87:13, v/v) in heat-resistant glass tubes on a heating block. The digests were
dissolved in 2.5% HNO3, and the concentrations of P, K, Ca, Mg, Cu, Zn, Fe, and Mn in the
digests were analyzed by Inductively Coupled Plasma-Mass Spectrometer (ICP-AES; Agilent
7500a, USA).
Data were analyzed using analysis of variance and treatment means were separated
using Duncan’s Multiple Range Test by SPSS 16.0 software.

RESULTS

Plant growth
No chlorosis was observed with both blueberries grown at pH 4.5. Only a mild
chlorosis occurred in newly formed leaves of ‘Climax’ at pH 5.3. Severe chlorosis appeared in
new leaves of both blueberries at pH 6.0, and bushes were small and stunted.
Plant height, stem-base diameter and shoot number were significantly inhibited in
‘Climax’ and A119 as the soil pH increased, and this inhibition effect was more pronounced
in ‘Climax’ than in A119 (Table 1). Compared to pH 4.5 treatment, plant height significantly
decreased 24 and 51% in ‘Climax’ and 17 and 52% in A119 at pH 5.3 and 6.0 treatment,
respectively. Meanwhile, stem-base diameter at pH 5.3 and 6.0 treatments decreased 13 and
39% in ‘Climax’ and 8 and 30% in A119, respectively. Compared to pH 4.5 treatment, shoot
number of both blueberries had no significant effects at pH 5.3 treatment, but significantly
decreased at pH 6.0 treatment in both blueberries.

Table 1. Effect of soil pH on plant height, stem-base diameter and shoot number of ‘Climax’
and A119.
Genotype pH Plant height (cm) Stem-base diameter (mm) Shoot number per plant
Climax 4.5 65.5±8.7 a1 9.57±0.52 a 5.8±2.1 a
5.3 49.8±4.8 b 8.34±0.67 b 4.8±1.0 ab
6.0 31.8±5.0 c 5.83±0.87 c 3.3±0.5 b
A119 4.5 64.8±3.5 a 10.94±1.29 a 5.0±1.4 a
5.3 53.8±3.4 b 10.01±0.17 a 4.3±0.5 a
6.0 30.8±3.6 c 7.64±0.51 b 2.8±0.5 b
1Different letters in the same column indicate significant difference between the treatments (P<0.05).
The dry weights of roots, stems, and leaves were markedly reduced with increasing
soil pH in both blueberries, whereas the root:shoot ratio was affected differently (Table 2).
The inhibition degree of dry weights of different organs was higher in ‘Climax’ than in A119
(Table 2). At the pH 5.3 treatment, dry weights of roots, stems, and leaves decreased 53, 41,
and 35% in ‘Climax’, and 28, 22, and 9% in A119, respectively, compared to pH 4.5
treatment. At the pH 6.0 treatment, dry weights of roots, stems, and leaves were reduced
approximately 80, 84, and 83% in ‘Climax’, and 71, 75, and 65%, respectively, compared to
pH 4.5 treatment. When the pH was raised from 4.5 to 6.0, root:shoot ratio of ‘Climax’
significantly increased firstly, and then decreased. No significant differences were found in
root:shoot ratio of A119 in different soil pH treatment.

271
Table 2. Effect of different soil pH on dry weight of different parts and root:shoot ratio of
‘Climax’ and A119.
Root dry weight Stem dry weight Leaves dry weight Root:shoot
Genotype pH
(mg plant-1) (mg plant-1) (mg plant-1) ratio
Climax 4.5 7.28±0.39 a 1 14.02±2.67 a 14.24±2.22 a 0.27±0.05 a
5.3 3.41±1.38 b 8.25±2.29 b 9.18±2.57 b 0.19±0.05 b
6.0 1.43±0.27 c 2.23±0.67 c 2.43±0.63 c 0.31±0.04 a
A119 4.5 12.00±1.88 a 15.27±1.13 a 10.86±0.49 a 0.46±0.06 a
5.3 8.60±1.49 b 11.89±1.52 b 9.91±1.34 a 0.40±0.04 a
6.0 3.50±0.42 c 3.81±0.70 c 3.81±0.69 b 0.47±0.08 a
1Different letters in the same column indicate significant difference between the treatments (P<0.05).

Leaf nutrition
The effect of soil pH on different leaf mineral nutrient content was varied (Tables 3
and 4). The levels of Fe and Mn of ‘Climax’ and A119 decreased as the soil pH increased,
whereas the levels of K, Ca and Mg increased. The Cu and Zn content were unaffected
irrespective of pH in both blueberries. The P content decreased in ‘Climax’ and remained
fairly constant in A119 as the soil pH levels increased.

Table 3. Effect of soil pH on leaf macronutrient content of ‘Climax’ and A119.

Genotype pH P (%) K (%) Ca (%) Mg (%)
Climax 4.5 0.11±0.01 a1 0.62±0.01 b 0.16±0.02 b 0.08±0.01 b
5.3 0.10±0.01 ab 0.62±0.04 b 0.22±0.04 a 0.08±0.01 b
6.0 0.08±0.02 b 0.76±0.12 a 0.26±0.04 a 0.09±0.01 a
A119 4.5 0.10±0.00 a 0.53±0.04 b 0.36±0.07 b 0.07±0.02 c
5.3 0.09±0.01 a 0.59±0.06 b 0.53±0.08 a 0.12±0.01 b
6.0 0.09±0.02 a 0.72±0.23 a 0.60±0.09 a 0.16±0.02 a
1Different letters in the same column indicate significant difference between the treatments (P<0.05).

Table 4. Effect of soil pH on leaf micronutrient content of ‘Climax’ and A119.

Genotype pH Cu (mg kg-1) Zn (mg kg-1) Fe (mg kg-1) Mn (mg kg-1)
Climax 4.5 4.04±1.22 a1 13.32±8.41 a 102.40±20.53 a 83.41±12.21 a
5.3 2.77±0.63 a 11.67±2.47 a 76.52±25.72 ab 11.06±3.61 b
6.0 2.67±0.50 a 6.08±1.42 a 32.23±6.00 b 6.07±1.38 b
A119 4.5 4.35±1.25 a 18.65±4.24 a 90.73±10.18 a 177.76±42.52 a
5.3 2.96±0.65 a 18.18±1.79 a 89.36±23.32 a 38.38±11.50 b
6.0 3.66±1.38 a 11.63±6.61 a 36.07±7.99 b 14.18±2.40 b
1Different letters in the same column indicate significant difference between the treatments (P<0.05).

Chlorophyll content and chlorophyll fluorescence

Chlorophyll content index (CCI), PSII maximum photochemical efficiency (Fv/Fm) and
effective light quantum yield [Y(II)] decreased as the soil pH increased in the two
blueberries (Table 5). Compared to pH 4.5 treatment, CCI and Fv/Fm significantly decreased
by around 48 and 3.6% in ‘Climax’, respectively, but they were not significantly decreased in
A119. Y(II) was unaffected in both blueberries at pH 5.3 treatment. At the pH 6.0 treatment,
CCI, Fv/Fm, and Y(II) were reduced approximately 76.19, 5.50, and 12.85% in ‘Climax’, and
53.02, 2.70, and 4.13% in A119, respectively, compared to the pH 4.5 treatment.

272
Table 5. Effect of different soil pH on chlorophyll content index, the maximal photochemical
efficiency of PSII (Fv/Fm) and the effective quantum yield of PSII[Y(II)]in leaves of
‘Climax’ and A119.
Genotype pH Chlorophyll content index Fv/Fm Y(II)
Climax 4.5 35.40±6.06 a1 0.787±0.014 a 0.685±0.029 a
5.3 18.40±9.43 b 0.759±0.013 b 0.653±0.010 a
6.0 8.43±0.46 b 0.746±0.004 b 0.597±0.031 b
A119 4.5 38.78±4.84 a 0.816±0.002 a 0.703±0.003 a
5.3 37.05±4.62 a 0.810±0.008 a 0.698±0.026 ab
6.0 17.83±7.46 b 0.794±0.009 b 0.674±0.011 b
1Different letters in the same column indicate significant difference between the treatments (P<0.05).

DISCUSSION
Blueberry plants grow best in acidic soil (pH 4.0-5.5). In previous research,
blueberries grew well in a low pH regime (pH 4.0), but poorly in the high pH regime (pH
6.5), and intermediately at the pH level of 4.5 (Finn et al., 1993). Plant weight also decreased
in the northern highbush blueberry cultivar ‘Bluecrop’ when the pH of nutrition solution
increased from 4 to 8, especially under low Fe or nitrate nitrogen conditions (Oertli, 1963).
Tsuda et al. (2014) found the rooting rates of highbush blueberry ‘Spartan’ was 0% and the
viability of roots was low, but the rooting rates of the hybrid clone JM4 between V.
corymbosum ‘Spartan’ and V. bracteatum was 91% and the viability of roots was high. These
results indicated high pH was not good for blueberry growth. However, most of the pH of soil
cannot reach this requirement, and is generally higher than this value. So the high value of
soil pH is the key factor limiting the blueberry growth and expansion.
In our research, we found the growth of two blueberries, A119, a southern highbush
blueberry selection, and ‘Climax’, a commonly used rabbiteye blueberry cultivar, was
significantly inhibited under the elevated soil pH, especially at pH 6.0 treatment, which
agreed with the results of previous studies (Finn et al., 1993; Oertli, 1963; Tsuda et al.,
2014). However, when the soil pH increased from 4.5 to 6.0, A119 had fewer declines in
most of the biomass indices than ‘Climax’, indicating that A119 had better adaptability to the
higher soil pH. Generally, the adaptability of southern highbush blueberry is lower than that
of rabbiteye blueberry, which has led to rabbiteye cultivars being the primary production in
southern China (He et al., 2009). In this study, the southern highbush blueberry selection
A119 was similar or even higher resistance to high pH than rabbiteye blueberry cultivar
‘Climax’.
One of the internal factors that caused poor growth of blueberries under higher pH is
nutrient imbalance in the tissue. Hall et al. (1964) found Ca, Mg, and Fe concentrations
increased or were unchanged, and P, K, and Mn concentrations and plant dry weight
decreased as pH level increased in lowbush blueberry. Haynes and Swift (1985) determined
that plant Mn, Zn, and Cu concentrations decreased and Fe concentrations first decreased
and then increased as soil pH level increased from 3.8 to 6.4 in highbush blueberry. Rosen et
al. (1990) found shoot K and Mn of ‘Northblue’ at pH 6.5 were significantly lower than that
at pH 4.5, and shoot N, P, Ca, Mg, Fe, Zn, and Cu concentrations were not affected between the
two pH levels. Shoot K, Fe, Mn, and Zn of ‘Searles’ (cranberry, V. macrocarpon) at pH 6.5 were
significantly lower than at pH 4.5, shoot Ca concentration was significantly higher, and shoot
N, P, Mg, and Cu concentrations were not affected between two pH conditions. In our
research, leaf K, Ca, and Mg concentrations increased, leaf Fe and Mn concentrations
decreased, and leaf Cu and Zn concentrations were unaffected when soil pH increased from
4.5 to 6.0, which indicated partial results consistent with the above reports. The present
results also agreed with previous reports about blueberry leaf chlorosis for iron deficiency
under high pH (Holmes, 1960). Holmes (1960) found that resolvable Fe content in solution
quickly decreased when solution pH increased. Haynes and Swift (1985) also found
extractable Fe content in soil decreased with increased soil pH. Rosen et al. (1990) suggested

273
most of the Fe appeared to be precipitated on the root surface as iron phosphate based on X-
ray microanalysis in cranberry, especially under high pH treatment. So the decrease of
blueberry Fe concentrations in leaves under high pH conditions may result from low Fe
resolution and precipitation as iron phosphate in root surface in this experiment.
High pH influences plant nutrient absorption, transport and distribution, leads to
nutrient imbalance, and further affects the growth and development of plants. For example,
iron deficiency can block the synthesis of chlorophyll, which leads to leaf chlorosis, and can
affect photosynthesis. Zhang et al. (2015) found that increase of rhizosphere pH decreased
chlorophyll concentrations and net photosynthesis of Pinus banksiana. We also observed
that the leaves of the two blueberries were chlorotic, and chlorophyll concentrations were
reduced when the soil pH raised, which could decrease photosynthetic rate and inhibit the
plant growth.
The change of chlorophyll fluorescence can indirectly reflect the change of plant
photosynthesis; thus we can be indirectly measure plant photosynthesis capacity by simple
determination of chlorophyll fluorescence. Some researchers have already used chlorophyll
fluorescence indicators to test the tolerance of plants to different stress, such as drought
resistance (Boureima et al., 2012), salt resistance (Rachoski et al., 2015), cold resistance
(Perks et al., 2004), and iron deficiency (Osó rio et al., 2014; Pestana et al., 2011). In this
study, we found that the chlorophyll fluorescence of the two blueberries declined with
increased soil pH. Moreover, chlorophyll fluorescence in A119, which was more tolerant to
high pH, decreased more slowly than in ‘Climax’, and illustrated that the chlorophyll
fluorescence index could also be used to evaluate blueberry tolerance to high soil pH.
In conclusion, the effect of different soil pH on plant growth, chlorophyll content,
chlorophyll fluorescence, and leaf mineral element content were studied in two blueberries,
A119, a southern highbush blueberry selection, and ‘Climax’, a rabbiteye blueberry cultivar.
We found plant height, stem-base diameter, shoot number, leaf dry weight, stem dry weight,
root dry weight, chlorophyll content index, PSII maximum photochemical efficiency and
effective light quantum yield showed decreasing trends with increased soil pH in the two
blueberries. A lower reduction was found in A119 than ‘Climax’, and indicated that A119 had
higher tolerance to high soil pH than ‘Climax’. Leaf Fe and Mn content of ‘Climax’ and A119
decreased as the soil pH increased, whereas K, Ca and Mg content increased. P content
decreased in ‘Climax’ and remained fairly constant in A119 as the soil pH levels increased.
Cu and Zn content were unaffected irrespective of pH in the two blueberries. The disorder of
leaf mineral element content could be a key factor to inhibit the growth of blueberry when
the soil pH was increased.

ACKNOWLEDGEMENTS
We gratefully acknowledge financial support from the opening fund of Jiangsu
province key laboratory for plant ex-situ conservation (QD201303).

Literature cited
Boureima, S., Oukarroum, A., Diouf, M., Cisse, N., and Van Damme, P. (2012). Screening for drought tolerance in
mutant germplasm of sesame (Sesamum indicum) probing by chlorophyll a fluorescence. Environ. Exp. Bot. 81,
37–43 http://dx.doi.org/10.1016/j.envexpbot.2012.02.015.
Finn, C.E., Luby, J.J., Rosen, C.J., and Ascher, P.D. (1993). Blueberry germplasm screening at several soil pH
regimes. I. Plant survival and growth. J. Am. Soc. Hortic. Sci. 118, 377–382.
Hall, I.V., Aalders, L.E., and Townsend, L.R. (1964). The effects of soil pH on the mineral composition and growth
of the lowbush blueberry. Can. J. Plant Sci. 44, 433–438 http://dx.doi.org/10.4141/cjps64-084.
Harmer, P.M. (1944). The effect of varying the reaction of organic soil on the growth and production of the
domesticated blueberry. Soil Sci. Soc. Am. Proc. 9 (C), 133–141 http://dx.doi.org/10.2136/sssaj1945.036159950
009000C0021x.
Haynes, R.J., and Swift, R.S. (1985). Effects of soil acidification on the chemical extractability of Fe, Mn, Zn and Cu
and the growth and micronutrient uptake of highbush blueberry plants. Plant Soil 84 (2), 201–212
http://dx.doi.org/10.1007/BF02143184.

274
He, S., Yu, H., and Gu, Y. (2009). Prospects and problems of blueberry growing in China. Acta Hortic. 810, 61–64
https://doi.org/10.17660/ActaHortic.2009.810.3.
Holmes, R.S. (1960). Effect of phosphorus and pH on iron chlorosis of the blueberry in water culture. Soil Sci. 90
(6), 374–379 http://dx.doi.org/10.1097/00010694-196012000-00010.
Oertli, J.J. (1963). Effect of form of nitrogen and pH on growth of blueberry plants. Agron. J. 55 (3), 305–307
http://dx.doi.org/10.2134/agronj1963.00021962005500030031x.
Osó rio, J., Osorio, M.L., Correia, P.J., de Varennes, A., and Pestana, M. (2014). Chlorophyll fluorescence imaging as a
tool to understand the impact of iron deficiency and resupply on photosynthetic performance of strawberry
plants. Sci. Hortic. (Amsterdam) 165, 148–155 http://dx.doi.org/10.1016/j.scienta.2013.10.042.
Perks, M.P., Osborne, B.A., and Mitchell, D.T. (2004). Rapid predictions of cold tolerance in Douglas-fir seedlings
using chlorophyll fluorescence after freezing. New For. 28 (1), 49–62 http://dx.doi.org/10.1023/B:NEFO.
0000031331.08847.49.
Pestana, M., Correia, P.J., David, M., Abadı́a, A., Abadı́a, J., and de Varennes, A. (2011). Response of five citrus
rootstocks to iron deficiency. J. Plant Nutr. Soil Sci. 174 (5), 837–846 http://dx.doi.org/10.1002/jpln.201000341.
Rachoski, M., Gazquez, A., Calzadilla, P., Bezus, R., Rodriguez, A., Ruiz, O., Menendez, A., and Maiale, S. (2015).
Chlorophyll fluorescence and lipid peroxidation changes in rice somaclonal lines subjected to salt stress. Acta
Physiol. Plant. 37 (6), 117 http://dx.doi.org/10.1007/s11738-015-1865-0.
Rosen, C.J., Allan, D.L., and Luby, J.J. (1990). Nitrogen form and solution pH influence growth and nutrition of two
Vaccinium clones. J. Am. Soc. Hortic. Sci. 115, 83–89.
Tsuda, H., Kunitake, H., Aoki, Y., Oyama, A., Tetsumura, T., Komatsu, H., and Yoshioka, K. (2014). Efficient in Vitro
screening for higher soil pH adaptability of intersectional hybrids in blueberry. HortScience 49, 141–144.
Yu, H., Gu, Y., Jiang, Y., and He, S. (2012). An update on blueberry growing in China. Int. J. Fruit Sci. 12 (1-3), 100–
105 http://dx.doi.org/10.1080/15538362.2011.619128.
Zhang, D., Yu, H., Jiang, Y., and Wang, C. (2009). Seedling selection of ‘Southmoon’ blueberry. Acta Hortic. 810, 65–
70 http://dx.doi.org/10.17660/ActaHortic.2009.810.4.
Zhang, W., Xu, F., and Zwiazek, J.J. (2015). Responses of jack pine (Pinus banksiana) seedlings to root zone pH and
calcium. Environ. Exp. Bot. 111, 32–41 http://dx.doi.org/10.1016/j.envexpbot.2014.11.001.

275


276