American Journal of Plant Sciences, 2010, 1, 47-54
doi:10.4236/ajps.2010.11007 Published Online September 2010 (http://www.SciRP.org/journal/ajps)
47
Characteristics of Gas Exchange in Three
Domesticated Anemone Species
Feihu Liu1*, Fei Li1, Xueni Liang2
1Faculty of Life Science, Yunnan University, Kunming, China; 2College of Adults and Professional Education, Yunnan University,
Kunming, China.
Received August 9th, 2010; revised August 27th, 2010; accepted August 31st, 2010
ABSTRACT
Seeds of three Anemone species were collected from the suburban areas of Kunming and planted in a nursery for three
and a half years at Kunming, Yunnan Province, China. Leaf gas exchange measurement indicated that these species
had similar one-peak diurnal trends of net photosynthetic rate (PN), although A. rivularis had lower transpiration rate
(TR), stomatal conductance (gs) and intercellular CO2 concentration (Ci), and higher stomatal limit in the afternoon.
Species differences in response of PN to photosynthetically active radiation (PAR) were observed, especially under
strong light. A. rivularis had the highest PN and Ci under strong light which correspon ded with its h ighest gs and TR. A.
rivularis had the highest light saturation point (LS P) (1000  mol m-2 s-1) and light compensa tion point (LCP) (69  mol
m-2 s-1), while A. hupehensis var. japonica had the lowest LSP (800
 mol m-2 s-1) and a lower LCP (53  mol m-2 s-1). But
the three species responded similarly to the change of CO2 concentration in the air from 0 to 350
 mol (CO2) mol-1, and
their observed CO2 compensation point showed little difference (47, 53 and 56
 mol (CO2) mol-1). Moreover, A. rivu-
laris had the highest apparent quantum yield (0.032), carboxylation efficiency (0.049), PN (11.68  mol (CO2) m-2 s-1)
and TR (5.36 mmol (H2O) m-2 s-1) based on the PN -PAR response. The results implied that A. rivularis is able to grow
well under higher radiation, while A. hupehensis var. japonica is the best one to grow under partial shade.
Keywords: CO2 Compensation Point, Gas Exchange, Light Compensation Point, Light Saturation Point
1. Introduction
Anemone is a newly developed species for cut flowers
that is also used as a potted plant. Many commercial Ane-
mone varieties have been released in other countries [1].
Anemone is an important genera (25 species, 4 subspe-
cies, and 9 varieties) in Yunnan province, China [2]. Inv-
estigations have been reported on taxonomy, cytology
and pollen morphology of Anemones [3, 4] and breeding
studies were carried out by Hoot [5], Jacob [6] and Lind-
ell [7]. However, photosynthesis has not been document-
ed for the genus.
Many studies are being done on development and uti-
lization of wild flower species for ornamental purpose, of
which cultivation of the wild species is one of the impor-
tant challenges [1]. Three wild Anemone species, collec-
ted from Yunnan Province, were successfully cultured in
a suburban plant nursery at Kunming, but they showed
different responses to the light environment. This was re-
asonable in view of their in situ growth environment, and
suggested the importance of controlling light intensity in
artificial culture of these wild plants. Consequently, these
Anemone species were presumed to have their own pecu-
liarities in photosynthesis. This paper aimed at studying
the photosynthetic characteristics of Anemone species and
their photosynthetic responses to the important compone-
nts of the environment, providing knowledge for control-
ling light conditions in the cultivation of Anemone spe-
cies.
2. Materials and Methods
2.1. Plants and Cultivation
Three Anemone representative species were selected acc-
ording to the distribution, leaf and plant form, and grow-
th habit. Seeds were collected from the suburban areas of
Kunming, Yunnan Province, China. A. vitifolia Buch.-
Han. Ex DC. (growing on a stony hillside at 2200 m a.s.l),
A. rivularis Buch.-Han. Ex DC. (growing on a grass hill-
side under sparse trees at 2230m a.s.l.) and A. hupehensis
Lemoine var. japonica (Thunb.) Bowles et Stearn (grow-
ing on a limestone wall at 2180 m a.s.l). Seedlings from

Characteristics of Gas Exchange in Three Domesticated Anemone Species
48
seeds planted in a suburban nursery of light loam soil wi-
th satisfied drainage at Kunming in early spring of 2000
(random block design with three replicates, at density of
20  40 cm in 3 m2 plots). The plants had grown under
light shade of trees (50%-100% of natural sunshine, var-
ied along with the angle of incidence), for three and a
half years at which time the tests were carried out.
Kunming (2501’E, 10241’N, 1896m a.s.l) has a dry
season from November through April and a wet season
from May through October, with a yearly mean temper-
ature of 14.9˚C, RH of 72%, and an annual rainfall of
1011 mm. The full natural sun radiation at the experime-
ntal site was recorded equivalent photosynthetically acti-
ve radiation 1500-2000 μmol m-2 s-1 on cloud-free days
throughout the growth season. The soil prior to the expe-
riment contained 150.3 mg kg-1 hydrolysable nitrogen,
25.1 mg kg-1 available phosphorus, 250.3 mg kg-1 availa-
ble potassium, 5% of organic matter and with a pH 7.45.
The plants were watered if necessary to avoid water
stress during the growing season and were fertilized tw-
ice a year before flowering, each time with 80 kg ha-1 of
urea (46% of N), 100 kg ha-1 of normal calcium superph-
osphate (18% of P2O5) and 65 kg ha-1 of potassium sulfa-
te (50% of K2O). The plants showed normal growing, fl-
owering, and fruit setting. The biomass was tested for the
species by harvesting all the plots when photosynthesis
measurement finished.
2.2. Photosynthesis Measurements and
Environmental Data
In late July, 2003 during flower bud stage, net photosyn-
thetic rate (PN) (3 plants per species, the last fully exp-
anded leaf per plant) in the three species was measured in
the field by a portable gas-exchange analyzer (LI-6400,
LICOR, Lincoln, NE, USA) (tested leaf area 6cm2, air
flow rate 400μmol s-1, stomatal ratio 0.5, a long tube was
used to draw the inlet air far from the operator in order to
minimize human impact on the CO2 levels). Stomatal co-
nductance (gs), intercellular CO2 concentration (Ci), tran-
spiration rate (TR), air temperature (Ta), ambient CO2
concentration (Ca), relative air humidity (RH) and photo-
synthetically active radiation (PAR) were also recorded.
Diurnal trend of PN was analyzed at 1-hour intervals
from 0700 to 1900 h on a cloud-free day. This was repea-
ted on one subsequent cloud-free day. Leaf water use eff-
iciency (LWUE) was calculated as PN / TR [8], and sto-
matal limitation (Ls) as (Ca - Ci) / Ca [9]. The environm-
ental conditions were recorded by the LI-6400 instrument
as shown in Figures 1(a), (b). Ca and RH did vary con-
siderably from early morning to nightfall.
The response of photosynthesis to light intensity was
measured. PN was tested from 0830-1130 h on a cloud-
free day respectively under PAR 2000, 1800, 1600, 1400,
1200, 1000, 800, 600, 400, 200, and 0 μmol m-2 s-1 (using
6400-02B light source) to set up a response of photosyn-
thesis to light intensity, when Ta, RH, and Ca were
22.5-28˚C, 45-65%, and 360-375 μmol (CO2) mol-1. On
one subsequent cloud-free day from 0830-1030 h, PN
was tested respectively under PAR 800, 600, 400, 200,
80, 40, and 0 μmol m-2 s-1 to determine accurate light
compensation points in different species, when Ta, RH,
and Ca were 23-26.5˚C, 50-65%, and 365-375 μmol
Figure 1. Diurnal trends of the photosynthetically active radiation (PAR), air temperature (Ta), ambient CO2 concentration
(Ca), relative humidity (RH), net photosynthetic rate (PN) and transpiration rate (TR) in the domesticated Anemone species.
The symbols and the vertical bars in the figure were mean ± standard deviation.

Characteristics of Gas Exchange in Three Domesticated Anemone Species
49
(CO2) mol-1. Sample leaves were first exposed to the hig-
hest PAR. When a constant PN was achieved PAR was
lowered step by step to total darkness. At each step, a
constant PN was achieved prior to recording the data. The
apparent quantum yield (AQY) was calculated as the
initial slope of the regression PN on PAR under condi-
tions of PAR 0, 40 and 80 μmol m-2 s-1 [10-12].
Light saturation points (LSP) were the turning point
areas on the curves in Figure 3(a), while light compen-
sation points (LCP) were derived from the linear correla-
tion of PN and PAR when PAR  80 μmol m-2 s-1 (Figure
4), here the constant dependent X (PAR) was considered
as LCP, i.e., the PAR value when PN = zero. CO2 compe-
nsation points (CCP) was determined by the linear corre-
lation between PN and Ca when Ca  100 μmol (CO2) mol-1
(Figure 5(a)), here the constant dependent X (CO2) was
considered as CCP, i.e., the CO2 value when PN = zero.
The response of photosynthesis to CO2 concentration
was measured from 0900-1100 h on another cloud-free
day, when Ta and RH were 24-27˚C and 55-65%. PAR
was set at 1200 μmol m-2 s-1 based on the PN-PAR re-
sponse (see Figure 3(a)), and Ca was set at 0, 50, 100,
150, 200, 250, 300, and 350 μmol (CO2) mol-1 by adjust-
ing the instrument. The carboxylation efficiency (CE)
was calculated as the initial slope of the regression PN on
Ci under conditions of Ca 0, 50 and 100 μmol (CO2) mol-1
[9,13].
2.3. Data Analysis
PN and TR values at the seven PAR settings i.e. 800, 1000,
1200, 1400, 1600, 1800 and 2000 μmol m-2 s-1 were used
for comparing PN and TR among species. Specific dif-
ference was determined by LSD test (Fisher) at  = 0.05.
Post hoc comparisons for the data in Table 1 and all dia-
grams were carried out using Statistica 5.0 (StatSoft Inc.
(1995) STATISTICA). The vertical bars in the figures
were means ± standard deviations (SD).
3. Results
3.1. Diurnal Patterns of PN and the Related Pa-
rameters
The three species had similar one-peak diurnal trends of
PN showing an asymmetric parabola-like curve (Figure
1(c)). A. vitifolia had a maximum PN at 1000 h, the other
two at 1100 h. At 1000, 1300-1500 and 1700-1800 h, A.
rivularis had lower PN values than A. vitifolia. TR
showed the same trend peaking at 1300 h (Figure 1(d)),
but A. rivularis had lower TR than the other two from
1200-1700 h.
The diurnal trends of gs in the three species were similar,
but species differences could be seen from the height of
peaks and curves, as well as the timing of peak value
(Figure 2(a)). Ci decreased up to 1000 h and increased
after 1700 h (Figure 2(b)), showing little change from
1000 to 1700 h at about 280 μmol (CO2) mol-1, with A.
rivularis being exceptionally low, the result from the
lower values of gs (Figure 2(a)).
LWUE peaked at 0900 h, decreasing from 0900 h to
1100 h due to TR increase (Figure 1(d)), then remaining
Figure 2. Diurnal trends of the stomatal conductance (gs), intercellular CO2 concentration (Ci), leaf water use efficiency
(LWUE, = PN / TR) and stomatal limitation (Ls, = (Ca - Ci) / Ca) in the domesticated Anemone species. The symbols and the
vertical bars in the figure were mean ± standard deviation.

Characteristics of Gas Exchange in Three Domesticated Anemone Species
50
Figure 3. Responses of the net photosynthetic rate (PN), intercellular CO2 concentration (Ci), transpiration rate (TR) and
stomatal conductance (gs) to photosynthetically active radiation (PAR) in the domesticated Anemone species. The symbols
and the vertical bars in the figure were mean ± standard deviation.
constant until 1800 h (Figure 2(c)). Ls showed diurnal
variations and all the species maintained high Ls values
from 1000 h to 1700 h (Figure 2(d)), but A. rivularis had
the highest values of Ls from 1200 – 1600 h.
3.2. Response of Photosynthesis to Light
Intensity
PN responses to PAR of the considered species showed
significant differences under the light intensity of 400
through to 2000 μmol m-2 s
-1. Under this condition, A.
rivularis had the highest PN, and A. hupehensis var. japo-
nica had the lowest, showing their difference in sensitiv-
ity to light intensity (Figure 3(a)). PN increased rapidly
when PAR increased from zero to 600 or 800 μmol m-2
s-1, although light depression of photosynthesis was not
observed within the short time (0900-1100 h) when PAR
did not exceed 1800 μmol m-2 s-1. LSPs were 1000 μmol
m-2 s-1 for A. rivularis and A. vitifolia, and 800 μmol m-2
s-1 for A. hupehensis var. japonica (Figure 3(a)). LCPs
of A. vitifolia, A. hupehensis var. japonica and A. rivu-
laris were 47  1.2 (SD), 53  1.4 and 69  1.6 μmol m-2
s-1 (Figure 4).
Ci responded to light intensity reversal to PN (Figure
3(b)) and reached 380-410 μmol (CO2) mol-1 when the
leaf was put in dark (PAR = 0). A. rivularis had higher Ci
values than the other two under light intensity of
200-2000 mol m-2 s
-1. Correlation analysis showed a
significantly negative correlation between PN and Ci (r =
–0.93*, data not shown). Moreover, Anemones showed a
stable TR that was influenced little by the increasing
PAR (Figure 3(c)), and A. rivullaris had the highest TR
values that corresponded with gs (Figure 3(d)).
3.3. Response of Photosynthesis to CO2
Concentration
To estimate CCP and CE, PN responses of the Anemone
species were measured under a range of Ca from 0 to 350
μmol (CO2) mol-1. PN increased when Ca increased from
04080200 400 600 800
PAR [  mol m
-2
s
-1
]
-4
0
4
8
12
PN [mol (CO2) m-2 s-1]
A. vitifolia
A. rivularis
A. hupehensis var. japonica
Figure 4. Photosynthetic rate (PN) of the Anemone species
under low photosynthetically active radiation (PAR) show-
ing the light compensation point. The symbols and the ver-
tical bars in the figure were mean ± standard deviation.

Characteristics of Gas Exchange in Three Domesticated Anemone Species
51
Figure 5. Responses of the net photosynthetic rate (PN) and
intercellular CO2 concentration (Ci) to ambient CO2 con-
centration (Ca) in the domesticated Anemone species. The
symbols and the vertical bars in the figure were mean±
standard deviation.
0 to 350 μmol (CO2) mol-1, showing insignificant differe-
nces among the considered species (Figure 5(a)). Ci sho-
wed same response to the change of Ca (Figure 5(b)).
CCP was 47  1.0 (SD) μmol (CO2) mol-1 for A. vitifolia,
53  1.1 for A. rivularis and 56  1.4 for A. hupehensis
var. japonica (Figure 5( a )).
3.4. Differences in PN, TR, CE, AQY and
Biomass
PN and TR were compared according to the data of PN /
TR-PAR responses within PAR 800-2000 μmol m-2 s-1 and
showed significant specific difference (Table 1). A.
rivularis had the highest PN and TR, while A. hupehensis
var. japonica had the lowest. The specific difference was
observed in biomass that showed a close correlation with
PN (r = 0.991*, Table 1).
CE varied from 0.036 to 0.049 among the species with
A. rivularis having the highest CE. The specific differ-
ence in CE showed a close correlation with PN values (r
= 0.999*, Table 1). AQY was in the range 0.023 to 0.032;
A. rivularis again had the highest value. AQY was also
closely associated with PN values (r = 0.999*, Table 1).
4. Discussion
4.1. Diurnal Trend of Photosynthetic Rate
In this experiment, the major environmental factors,
PAR, Ta and air RH changed remarkably during the day as
shown in Figures 1(a), (b). This situation is normal at
our experimental site and is the driver of PN daily change.
No increase of PN was observed in the afternoon while a
midday depression in photosynthesis was found in
Anemone species collected from the alpine region in
suburban areas of Kunming (1896 m a.s.l.). PN increased
before 1000 or 1100 h and decreased from 1000 or 1100
h (dependent on species) onwards. The increase of PN in
the morning was driven by the increase of PAR and gs (Fig-
ures 1(a), (c); Figure 2(a)), and decrease of PN from the
late morning to the afternoon was caused by the decrease of
gs (Figure 1(c); Figure 2(a)). A close link between PN
and gs and/or PAR (only in the morning) was observed in
the investigation of the diurnal trends of photosynthetic
rate for Ginkgo biloba, Paspalum notatum, and Enkleia
malaccensis [14-16]. Whereas gs decrease in this ex-
periment resulted from the integrative action of adverse
ecological factors such as the rise of Ta (> 30˚C), de-
crease of air RH (< 35%) and Ca (< 386 μmol (CO2) mol-1),
and strong light (> 1500 μmol m-2 s-1) (Figures 1(a), (b)).
A strong light of no more than 1800 μmol m-2 s-1 did not
depress the photosynthesis of the Anemones in the field
(Figure 3(a)). This may reflect the intrinsic adaptation of
the Anemone species to the mild ecological environments
in their native habitats of moderate Ta, higher air RH but
strong light. The decrease of gs from 1000 or 1100 h on-
wards and the decrease of TR in the afternoon were
caused by the increase of stomatal limitation (Ls)
Table 1. Carboxylation efficiency (CE), apparent quantum yield (AQY), Photosynthetic rate (PN), transpiration rate (TR)
and biomass of the Anemone species.
Taxon CE AQY PN TR Biomass (kg m-2)
A. rivularis 0.0491  0.0014a 0.0317  0.0005a 11.68  0.448a 5.36  0.354a 1.02  0.055a
A. vitifolia 0.0417  0.0011b 0.0275  0.0007b 9.25  0.346b 3.22  0.351b 0.83  0.071b
A. hupehensis var. japonica 0.0355  0.0013c 0.0229  0.0009c 8.05  0.420c 2.96  0.372b 0.78  0.069b
PN and TR values at the seven PAR settings i.e. 800, 1000, 1200, 1400, 1600, 1800 and 2000μmol m-2 s-1 were used for comparing PN and TR among species.
Numbers in the table are means  SD. Same letters show no statistical difference among species by LSD test (Fisher) at  = 0.05. Correlation coefficients of PN
–AQY and PN–Biomass were 0.999* and 0.991*.

Characteristics of Gas Exchange in Three Domesticated Anemone Species
52
(Figure 1(d); Figures 2(a), (d)). Here TR seemed to
have a delayed response to the change of Ls, but in fact
TR was tightly linked to the increasing Ta, PAR and va-
por pressure deficit based on leaf temperature (VpdL),
and decreasing RH (Correlation coefficients of TR with
PAR, Ta, VpdL and RH were derived as 0.89*, 0.86*,
0.61* and –0.72*, respectively) (data not shown). This
led to a high level of TR around noon even though at this
time gs decreased and/or Ls increased. Therefore, PAR, Ta
and RH were the major drivers of TR (Figures 1(a), (b)
and (d)), as VpdL negatively correlated with RH (r =
–0.94*) (data not shown).
The decrease of Ci is the major criterion for verifying
Ls if PN decreased or stayed at a low level [17]. In our
experiment, Ci obviously decreased in the morning be-
fore 1100 h that reflected the normal changes from night
(where Ci increased over time because of respiration) to
daytime from increased intercellular CO2 use resulting
from increasing photosynthesis. The decrease of Ci in A.
rivularis from 1100 h until 1600 h (Figure 2(b)) was
likely caused by the decrease of gs (Figure 2(a)) since PN
decreased as well during this period (Figure 1(c)) and
water stress was avoided by watering. Based on the
above argument, the decrease of PN from the late morn-
ing to the afternoon was mainly caused by a shortage of
CO2 resulted from an increase of Ls and decrease of gs
(Figure 1(c); Figures 2(a), (d)). It is well known that the
increase of Ci is the reason for non-stomatal limitation of
photosynthesis when PN is at a low level [18]. Therefore,
the rapid decrease of PN after 1700 h was caused by an
increase of non-stomatal limitation (e.g., carboxylation
resistance) that quite likely resulted from the decrease of
PAR (Figure 1(a)), for even if gs was at a low level, Ci
increased and Ls decreased rapidly during this time.
There is a strong correlation between PN and gs because
stomata respond to the changes in assimilation via Ci.
Correlation analysis revealed a significantly positive
correlation between PN and gs (r = 0.77*), and a negative
correlation between PN and Ci (r = –0.73*) (data not
shown).
When PAR was set to zero, the PN values of –2.5- –2.0
μmol (CO2) m-2 s-1 might reflected the respiration rates of
the Anemones under the experimental conditions (Figure
4).
4.2. Response of Photosynthesis to Light
Intensity
In lettuce [19], tobacco [20], and Phaseolus vulgaris [21],
PN along with the increase of light intensity increased to
a peak, then decreased if the light intensity increased
further. However, in ramie (Boehmeria nivea (L.) Gaud.)
[22], tomato [23], cucumber [17], and cotton [24], as lig-
ht intensity increased, PN increased and kept on at a high
level within a wide range of light intensity. The depres-
sion of photosynthesis in tobacco when PAR > 600 μmol
m-2 s-1 was due to the reduction in activities of PSI and
PSII, carbonic anhydrase and electron transfer speed [20].
A large decrease in stomatal conductance to water va-
pour in Phaseolus vulgaris leaves exposed to strong light
was found [21].
In our experiments, strong light of no more than PAR
1800 μmol m-2 s-1 did not depress photosynthesis, althou-
gh differences were observed in utilization of light ene-
rgy under high light intensity (Figure 3(a)) with ade-
quate soil water, Ta 23-30˚C and RH 45-65%. Further-
more, the sample leaf was put under a strong light for a
few minutes for achieving a constant PN before going to
a lower PAR. Therefore, it needs more investigations to
uncover whether or not the Anemone species have a
photosystem insensitive to the change of light intensity.
The experiment results, based on single leaf test, sug-
gested that PAR 800-1000 μmol m-2 s
-1 is enough for a
satisfactory PN and meets the needs for photosynthetic
products for normal growth and development of Anem-
one cultivation.
Among the three species, A. rivularis was from a grass
hillside under sparse trees with good sunshine and had a
different response to light from other species, showing a
high LCP and LSP. Whereas, A. hupehensis var. japonica
was from a limestone wall with only intermittent sunshi-
ne and is probably less well adapted to continuous strong
light. The light response data, low LCP and LSP, and low
maximum PN value all indicated that this species is the
best one to grow under partial shade.
4.3. Response of Photosynthesis to CO2
Among PN, TR, gs, and Ci, only PN and Ci were observed
to increase regularly as Ca increased, and the response of
PN and Ci to the change of Ca showed little difference am-
ong the species (Figures 5(a), (b)). When Ca was set to
zero, PN was –3 μmol (CO2) m-2 s-1 (Figure 5(a)) which
was roughly considered as the sum of respiration and
photorespiration under the experimental conditions. This
was unlike the PN values at PAR = 0 μmol m-2 s-1 where
the minus PN values just presented the respiration rates.
The CO2 saturation curve of photosynthesis in the con-
sidered species is still in need of study.
A higher CCP (60-75 μmol (CO2) mol-1), but a much
lower LCP (13-28 μmol m-2 s
-1), and similar diurnal
curves of PN and TR were observed in domesticated Pri-
mula species collected from an alpine area of north-wes-
tern Yunnan Province, China [25]. This reflected the si-
milarity in photosynthetic daily patterns of Anemones and
Primulas as alpine plants, and the dissimilarity in light and
CO2 utilization qualities of plants of the two genera, which
should be taken into consideration for their cultivation.

Characteristics of Gas Exchange in Three Domesticated Anemone Species
53
5. Conclusions
The Anemone species described here showed similar one-
peak diurnal trends of PN, although A. rivularis had lo-
wer TR, gs, and Ci, and higher Ls in the afternoon. Spec-
ific differences in response of PN to PAR were observed,
especially under strong light. A. rivularis had the highest
PN and Ci under strong light which corresponded with its
high gs and TR. A. rivularis had the highest LSP (1000
mol m-2 s-1) and LCP (69 mol m-2 s-1), while A. hupe-
hensis var. japonica had the lowest LSP (800 mol m-2
s-1) and a lower LCP (53 mol m-2 s-1). All these species
responded similarly to the change of Ca from 0 to 350
mol (CO2) mol-1, and their CCP showed little difference.
Moreover, A. rivularis had the highest AQY, CE, PN and
TR based on the PN-PAR response. In conclusion, A.
rivularis is able to grow well under higher radiation,
while A. hupehensis var. japonica is the best one to grow
under partial shade.
6. Acknowledgements
The authors are grateful to the reading and comments
given by Gordon Rowland, University of Saskatchewan.
Thanks are expressed to Su WH of Yunnan University
for his kind help in the equipment operation.
REFERENCES
[1] Y. L. Yuan, J. L. Zhao and W. P. XU, “Anemone of Mor-
phology Characteristic and Introduction Apply Study,”
Acta Botanica Boreali-Occidentalia Sinica, Vol. 17, No.
5, 1997, pp. 134-136.
[2] Botanical Institute of Kunming CAoS, “Flora Yunna-
nica,” Science Press, Beijing, Vol. 11, 2000, pp. 183-204.
[3] M. Y. Fang and M. Y. Yang, “A Study on Pollen Morph-
ology and Evolution of the Genus Anemone from Sichu-
an,” Journal of Sichuan University, Vol. 31, No. 2, 1994,
pp. 246-258.
[4] Q. E. Yang, “Cytology of Ten Species in Anemone, One
in Anemoclema and Six in Clematis (Trib. Anemoneae,
Ranunc ulace ae) form China,” Acta Phytotaxonomica Sinica,
Vol. 40, No. 5, 2002, pp. 396-405.
[5] S. B. Hoot, “Phylogenetic Relationships in Anemone (Ranu-
culaceae) Based on Morphology and Chloroplast DNA,”
Systematic Botany, Vol. 19, No. 1, 1994, pp. 169-200.
[6] Y. Jacob, “Breeding of Anemone Coronaria Tetraploid
Hybrids,” Proceedings of the Seventh International Sym-
posium on Flower Bulbs, Vol. II, Herzliya, Israel. Acta
Horticulturae, Vol. 430, 1997, pp. 503-508.
[7] T. Lindell, “Breeding Systems and Crossing Experiments
in Anemone Patens and in the Anemone Pulsatilla Group
(Ranunculaceae),” Nordic Journal of Botany, Vol. 18, No.
5, 1998, pp. 549-561.
[8] J. J. Vacher, “Responses of Two Main Andean Crops,
Quinoa (Chenopodium quinoa Willd) and Papa Amarga
(Solanum juzepczukii Buk.) to Drought on the Bolivian
Altiplano: Significance of Local Adaptation,” Agriculture,
Ecosystems & Environment, Vol. 68, No. 1-2, 1998, pp.
99-108.
[9] G. D. Farquhar and T. D. Sharkey, “Stomatal Conduc-
tance and Photosynthesis,” Annual Review of Plant Phys-
iology, Vol. 33, 1982, pp. 317-345.
[10] J. Ehleringer and O. Björkman, “Quantum Yields for CO2
Uptake in C3 and C4 Plants,” Plant Physiology, Vol. 59,
No. 1, 1977, pp. 86-90.
[11] J. Ehleringer and R. W. Pearcy, “Variation in Quantum
Yield for CO2 Uptake among C3 and C4 Plants,” Plant
Physiology, Vol. 73, No. 3, 1983, pp. 555-559.
[12] Z. Y. Tao and Q. Zou, “Effect of Strong Irradiance and
Shortly Elevated CO2 Concentrations on Photosynthetic
Efficiencies in Maize and Soybean Leaves,” Acta Botanica
Boreali-Occidentalia Sinica, Vol. 25, No. 2, 2005, pp.
244-249.
[13] Z. J. Wang, T. C. Guo, Y. J. Zhu, Y. H. Wang, J. H.
Wang and M. Zhao, “Comparison of CO2 Assimilation
Capacity in Flag Leaf for Super High Yield Wheat with
Different Spike Type,” Acta Agronomica Sinica, Vol. 30,
No. 8, 2004, pp. 739-744.
[14] S. Pandey, S. Kumar and P. K. Nagar, “Photosynthetic
Performance of Ginkgo Biloba L. Grown under High and
Low Irradiance,” Photosynthetica, Vol. 41, No. 4, 2003,
pp. 505- 511.
[15] R. V. Ribeiro, G. B. Lyra, A. V. Santiago, A. R. Pereira,
E. C. Machado and R. F. Oliveira, “Diurnal and Seasonal
Patterns of Leaf Gas Exchange in Bahiagrass (Paspalum
Notatum Flugge) Growing in A Subtropical Climate,”
Grass and Forage Science, Vol. 61, No. 3, 2006, pp. 293
-303.
[16] A. C. Tay, A. M. Abdullah, M. Awang and A. Furukawa,
“Midday Depression of Photosynthesis in Enkleia Ma-
laccensis, a Woody Climber in a Tropical Rainforest,”
Photosynthetica, Vol. 45, No. 2, 2007, pp. 189-193.
[17] D. H. Ma, J. A. Pang, Z. R. He and S. J. Li, “Effect of
Environmental Factors on the Photosynthetic Characteris-
tics of Cucumber Seedlings,” Acta Botanica Boreali-Occ-
identalia Sinica, Vol. 12, No. 4, 1997, pp. 97-100.
[18] D. Q. Xu, “Some Problems in Stomatal Limitation Analy-
sis of Photosynthesis,” Plant Physiology Communications,
Vol. 33, No. 4, 1997, pp. 241-244.
[19] P. P. Li, Y. G. Hu, Y. G. Zhao, X. J. Ying and H. P. Mao,
“Comprehensive Model on the Effect of CO2 Enrichment
on Lettuce Photosynthesis in Greenhouse,” Transaction
of the CSAE, Vol. 17, No. 3, 2001, pp. 75-79.
[20] L. Jiang, S. Q. Cao, X. Dai and X. M. Xu, “Effect of Dif-
ferent Light Intensity on Photosynthesis of Tobacco,”
Journal of Chinese Tobacco, Vol. 6, No. 4, 2000, pp.
17-20.
[21] L. Guidi, M. Tonini and G. F. Soldatini, “Effects of High
Light and Ozone Fumigation on Photosynthesis in Phase-
olus Vulgaris,” Plant Physiology and Biochemistry, Vol.
38, No. 9, 2000, pp. 717-725.

Characteristics of Gas Exchange in Three Domesticated Anemone Species
54
[22] Q. Q. Guo, “Study on the Photosynthetic Characteristics
of Ramie Leaves in Different Varieties and Their Rela-
tionship with Yield Formation of Ramie. II. The Physiol-
Ecological Characteristics of Leaf in Photosynthesis,”
Journal of Hunan Agricultural College, Vol. 17, No. 3,
1993, pp. 76-79.
[23] C. Stanghellini and J. A. Bunce, “Response of Photosyn-
thesis and Conductance to Light, CO2, Temperature and
Humidity in Tomato Plants Acclimated to Ambient and
Elevated CO2,” Photosynthetica, Vol. 29, 1993, pp. 487
-497.
[24] H. Z. Dong, W. J. Li, W. Tang and Z. H. Li, “Photosyn-
thetic Characteristics of Field Grown Cotton Leaves,” Sh-
angdong Agricultural Science, No. 6, 2000, pp. 7-10.
[25] F. H. Liu, S. M. Hou and X. N. Liang, “Gas Exchange
Characteristics of Four Domesticated Primula Species,”
New Zealand Journal of Crop and Horticultural Science,
Vol. 34, 2006, pp. 403-411.