Study on the Influence of Three Production Methods on Blueberry

Photosynthesis
Yadong Lia, Chen Wei, Zhang Zhidong, Liuhaiguang and Wulin
Department of Horticutlure,
Jilin Agricultural University,
Changchun, 130118
P.R.China

Key words: blueberry, ‘bluecrop’, culturing pattern, photosynthetic characteristic

Abstract
The influence of three production methods (open-field, glasshouse, plastic
house) on photosynthetic parameters of Vaccinium corymbosum ‘Bluecrop’ were
compared, including net photosynthetic rate (Pn), stomatal conductance (Gs), and
intercellular CO2 concentration (Ci). For all production methods the diurnal curve
for ‘Bluecrop’ Pn had two peaks and a midday depression of Pn. The minimum Gs
under the three production methods occurred at 14:00 and the time of maximum Gs
varied with production method. The maximum Gs for plants grown in an open-field
occurred at 08:00 and 18:00 and the maximum Gs for plants grown in the
glasshouse and plastic house occurred at 10:00 and 16:00. The Ci in plants in all
production methods decreased between 06:00 and 10:00, remained low between
10:00 and 16:00, and increased after 16:00. The decline in Pn between 10:00 and
14:00 contributed to stomatal-limitation, while there was no stomatal limitation to
Pn at any other time during the day.

INTRODUCTION
Leaf photosynthesis is one of the most important factors affecting plant
development and yield and it is estimated that about 90% of fruit trees dry matter comes
from leaf photosynthesis (Massai et al., 2004; Su Huairui, 1993). Several authors describe
leaf photosynthesis on fruit trees; however, most reports concentrate on diurnal trends in
photosynthesis on apple, citrus, and grape (Chen and Zhang, 1993; Cheng et al., 1992;
Cao, 1997). There are several reports describing leaf photosynthesis on blueberry plants
but the studies do not report consistent or extensive results (Anderson, 1989; Forsyth and
Hall, 1965; Moon et al., 1987; Li et al., 1998, 1995; Zhang et al., 1999).
Fruit production in greenhouses has become a popular production method to
increase fruit production profits in China. Compared with open-field production,
greenhouse production systems significantly decrease solar radiation and can affect
photosynthesis, plant growth and fruit development. The objective of our study was to
describe diurnal changes in leaf photosynthesis of ‘Bluecrop’ grown in open-field and
greenhouse production methods. The results will improve our basic knowledge of
potential photosynthesis differences between plants grown by the methods.

MATERIALS AND METHODS

The experiment was conducted in the blueberry experimental farm of the Jilin
Agricultural University (43°88'N, 125°35'E). The growing substrate was a 1:1:1 (v/v) mix
of a black soil type (pH 7.0, 1% organic matterpeat and organic fertilizer. The pH of the
substrate was adjusted to 5.5 using 2 kg sulfur/m3. Three-3-year-old ‘Bluecrop’ plants
were planted in the open-field, glasshouse, or plastic house in April 2001.
In 2004, plants grown by each method (glasshouse, plastic house, and open-field)
were selected to determine leaf photosynthetic characteristics. Five plants per plot and
five plants per treatment were randomly selected for measurement during the stage of fast
shoot growth early in the growing season. This stage occurred in early May on plants in
a
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the glasshouse, early June for plants in the plastic house and late June for plants in the
open-field).
All measurements were taken on days with clear skies. On each of the plants three
functional leaves on the outside of the canopy were selected to measure Pn (net
photosynthetic rate), Gs (stomatal conductance), Ci (intercellular CO2 concentration) and
environmental factors such as leaf temperature, air temperature, air CO2 concentration and
PAR using a TPS-1 (PP Systems, made in U.K.). Measurements were taken every two
hours between 06:00 and 18:00). Stomatal limitation (Ls) was calculated as described by
Berry (1982): Ls=1-Ci/(C0-Г); where Ci= intercellular CO2 concentration; C0 = air CO2
concentration around leaf; and Г = CO2 compensation point.

RESULTS

Diurnal Variation in Net Photosynthetic Rate (Pn)

In all three production methods, diurnal variation in Pn ‘Bluecrop’ leaves was a
two-peak curve (Fig. 1A). The first and highest leaf Pn occurred at 06:00 on plants grown
in the open-field (late June measurement) and occurred at 08:00 on plants grown in the
glasshouse (early May measurement) and plastic house (early June measurement). In all
three production methods, leaf Pn declined with the increasing PAR and air temperature
during the day and reached a minimum at midday (midday photosynthetic depression).
When plants were grown in the glasshouse and plastic house the midday photosynthetic
depression occurred at 12:00 and a second peak in Pn occurred at 14:00. When plants
were grown in the open field the midday photosynthetic depression occurred at 14:00 and
a second peak in Pn occurred at 16:00. In all production methods, after the second peak in
leaf Pn in the afternoon, Pn declined rapidly with the decreasing of PAR and reached a
second minimum at 18:00.
The rates of Pn and time of maximum and minimum Pn values can not be directly
compared between production methods because measurements were taken at different
times of the year; however in general Pn was the highest for plants grown in the open-
field (late June), and the lowest for plants grown in the glasshouse (early May). Daily Pn
peak for plants in the open-field, plastic house and glasshouse were 9.64 µmol
CO2·m-2·s-1, 8.8 µmol CO2·m-2·s-1 and 6.95 µmol CO2·m-2·s-1, respectively.
Diurnal Variation in Leaf Stomatal Conductance (Gs)
In the glasshouse and the plastic house, diurnal variation in Gs was two-peak
curve with peaks of similar sizes (Fig. 1B). The first peak in GS in plants grown in the
glasshouse and plastic house occurred at 10:00 followed by a decline in Gs to a minimum
at 14:00. The second peak in GS in plants grown in the glasshouse and plastic house
occurred at 16:00. In the open-field, diurnal variation in Gs was also a two-peak curve but
peaks were not similar in size (Fig. 1B). The first and highest peak in GS in open-field
plants occurred at 08:00 followed by a decline in Gs to a minimum at 14:00. The second
and smaller peak in Gs in open-field plants occurred at 18:00.

Diurnal Variation in Intracellular Carbon Dioxide Concentration (Ci)

Diurnal variation in leaf Ci was a reverse parabola that was similar for plants
grown by different methods (Fig. 1C) and similar as air CO2 concentrations measured at
the same time (data not presented). For all production methods, leaf Ci declined between
06:00 and 12:00 and increased from 12:00 to 18:00.

Diurnal Variation in Photosynthetically Active Radiation (PAR)

In the glasshouse, PAR increased between 06:00 and 10:00 then decreased (Fig.
1D) (early May measurement). In the plastic house and open-field PAR increased
between 06:00 and 12:00 then decreased (Fig. 1D) (June measurements).

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Diurnal Variation in Leaf Stomatal Limitation (Ls) and the Pn/Gs Ratio
Diurnal variation in leaf stomatal limitation (Ls) was similar for plants in all
production methods (Fig. 1E). The Ls declined between 06:00 and 10:00 corresponding to
the time when there was a rapid decline in PAR (Fig. 1D). The Ls was relatively level
between 10:00 and 16:00 and increased rapidly after 16:00.
In the glasshouse and plastic house, leaf Pn/Gs decreased between 06:00 and
18:00. Leaf Pn/Gs changed dramatically in field grown plants throughout the day. The
highest leaf Pn/Gs occurred at 14:00 and the lowest at 16:00

DISCUSSION
High temperature or water stress is known to decrease stomatal conductance
resulting in a decrease in Ci and leaf Pn. The midday photosynthetic depression that
occurs in blueberry leaves is indicative of this response. Xu and Shen (1997) separated a
decrease in plant photosynthesis into two factors, a stomatal factor (stomatal closing) and
non-stomatal (leaf cell activity decreasing) factor (Liu and Guo, 1998). Farquhar et al.
(1982) used Ls to determine if a decrease in photosynthesis is a result of a stomatal factor
or non-stomatal factor when leaf Pn and Ci tended to decease. That is, stomatal closing
can be considered the main reason for a decrease in leaf photosynthesis if Ls increases.
Mediavilla et al. (2002) suggested using Pn/Gs as a standard index to determine whether a
stomatal factor or non-stomatal factor is responsible for a decrease in leaf photosynthesis.
According to Mediavilla’s standard index, if stomatal closing is the main reason for a
decrease in leaf photosynthesis, Pn/Gs should increase with increasing Ci.
In our study, leaf Pn, Ci and Ls of ‘Bluecrop’ grown in the open field decreased
but Pn/Gs increased from 06:00 to 08:00. This indicates that the decrease in Pn at this
time was caused by stomatal factors and non-stomatal factors. Leaf Pn, Ci, Ls and Pn/Gs
of plants grown in the open field also decreased from 08:00 to 10:00, indicating the
decrease in Pn is caused by non-stomatal factors. Between 10:00 and 14:00 and 16:00 and
18:00, leaf Pn of plants grown in the open field decreased but leaf Ci did not decrease.
Using Pn/Gs as the standard index during these times the decrease in Pn/Gs indicates
stomatal factors are the main reason for the decrease in leaf Pn. Leaf Pn and Ci variation
in plants grown in the glasshouse and plastic house were similar as that of plants in the
open field. Between 08:00 and 10:00 Pn of plants in the greenhouse and plastic house was
limited by non-stomatal factors and between 10:00 and 12:00 and 14:00 and 18:00 Pn was
limited by non-stomatal factors. Under the measurement conditions in our experiment,
leaf Pn in plants grown in the glasshouse and plastic house were limited by non-stomatal
factors and leaf Pn in plants grown in the open field was limited by non-stomatal factors
and stomatal factors.
Many researchers indicate light intensity, temperature and water are the main
factors limiting to leaf photosynthesis. According to our study, the highest PAR occurred
at 10:00 for plants grown in the open field and at 12:00 for plants grown in the glasshouse
and plastic house. This indicates that light intensity may be the main reason for leaf
midday photosynthetic depression in blueberry. Our data also indicate leaf temperature on
plants grown in the glasshouse reached a maximum at 10:00 (34.0°C) and remained
relatively high until 12:00. Leaf temperature reached a maximum at 14:00 for plants
grown in the plastic house and the open field (33.4°C and 33.5°C, respectively). The
optimal temperature for Rubisco activity is 25–30°C (Xu and Shen, 1997), higher
temperatures significantly limits Rubisco activity and decreases leaf Pn. Higher
temperatures can also increase plant respiration that may decrease leaf Pn ((Salvucci,
1989). Our results suggest the high temperature in the glasshouse and plastic house may
be responsible for decreasing blueberry leaf Pn at certain times of the year.

CONCLUSION
Light is a key factor in plant photosynthesis. There was a significantly difference
in PAR between glasshouse, plastic house and open field production methods for
blueberry. These differences, while partially a function of the time of year measurements

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were taken, also are reflective of how these different production methods may influence
leaf photosynthesis. While seasonal variation may be partially accountable for differences
in the magnitude of Pn daily variation of ‘Bluecrop’ leaf Pn was two-peak curve in all
production systems and midday photosynthetic depression occurred in all production
systems.

ACKNOWLEDGEMENTS
This research was financially supported by Nonprofit Industry Special Scientific
Funds (nyhzx07-028 and 2006-G25).

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Figures

12 140
greenhouse greenhouse
A plast ic house B plastic house
open field open field
10 120

Leal Gs (mmol H 2 O 2·m -2 ·s -1)

Leaf Pn (µmol CO 2·m -2 ·s -1)

100
8
80
6
60
4
40

2
20

0 0
6 8 10 12 14 16 18 6 8 10 12 14 16 18
Tim e of day (h ) Tim e of day (h )
1000 1400
C greenhouse greenhouse
plastic house D plastic house
900 open field 1200 open field
800
-1 )

1000
2 ·mol

700
·s -1 )

600
-2

800
PAR (µmol·m
Leaf Ci (µmolCO

500
600
400
300 400
200
200
100
0 0
6 8 10 12 14 16 18 6 8 10 12 14 16 18
Time of day (h) Time of day (h)
3 1. 2
E greenhouse F greenhouse
plastic house plastic house
2. 5 open field open field
1
2
-1 )

0. 8
1. 5
Pn/Gs (mmol·mol

1
Ls

0. 6

0. 5
0. 4
0
0. 2
- 0. 5

-1 0
6 8 10 12 14 16 18 6 8 10 12 14 16 18
Time of day (h) Time of day (h)

Fig. 1. Diurnal changes in (A) leaf net photosynthesis (Pn), (B) leaf stomatal conductance
(Gs), (C) leaf internal carbon dioxide concentration (Ci), (D) photosynthetically
active radiation (PAR), (E) stomatal limitation (Ls), and (F) Pn/Gs ratio of 3-year-
old ‘Bluecrop’ blueberry plants grown in a glasshouse, a plastic house, or an open
field.

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