J . Amer. Soc. Hort. Sci. 108(4):541-544. 1983.
Effect of Shade on Photosynthesis, Specific Leaf
Weight, Leaf Chlorophyll Content, and
Morphology of Young Peach Trees
Frank Kappel1 and J.A. Flore
Department of Horticulture, Michigan State University, East Lansing, MI 48824
Additional index words. Prunus persica, branch angle
Abstract. One-year-old peach [Prunus persica (L.) Batsch cv. Redhaven] trees were grown outdoors under 4 different
shade levels (100%, 36%, 21 %, and 9% of full sun). Shoot length, internode length, and node number were unaffected
by shade, but heavy shade (9% full sun) reduced stem diameter 24%. Compared to those in full sun, average leaf
areas were increased 18%, 30%, and 20% by 36%, 21%, and 9% shade, respectively. Shading caused a more
horizontal leaf orientation and lower specific leaf weight. Both average and maximum photosynthetic rate per unit
area or per mg chlorophyll decreased with shading, but increased with shading when expressed per unit dry weight.
Chlorophyll content per unit leaf area increased with shading, but stomatal resistance was unaffected. Leaves from
shade treatments became light-saturated between 400 and 600 |xE s *111 L; full-sun leaves became light-saturated at
700-900 *jiE s _Im”2.
The absorption and utilization of photosynthetically active achieved by growing trees outside without shading. Solar ra
radiation (PAR) is one of the most important factors that limits diation was reduced with pipe frame structures (3.7 x 2.4 x
plant productivity (11, 12). Shading reduces flower bud initia 1.8 m) covered with black polypropylene shade fabric (A.H.
tion, fruit size, fruit quality and color, number of new shoots, Hummert Co., St. Louis, Mo.), which transmitted an average
and plant fresh weight (5, 9, 12, 18, 19). Tree size, spacing, of 36%, 21%, or 9% PAR (measured with a LI-COR LI-190S
and training system have a dramatic effect on light conditions quantum sensor connected to a Model LI 188 quantum/radi-
within the canopy (5, 8, 12). ometer/photometer). Ventilation prevented temperature differ
The photosynthetic characteristics of an individual leaf or ences of greater than ± 3°C and relative humidity differences
plant can be greatly influenced by PAR (3, 4). Plants grown in greater than ±5% . Spectral radiometer (ISCO Model SR port
the shade have lower maximum photosynthetic rates than those able spectroradiometer, Lincoln, Neb.) determinations con
grown in full sun (4), but are often more efficient at low PAR firmed that all wavelengths in the 380-750 nm range were reduced
levels. Barden (2) found that for apple, net photosynthesis, dark equally. Four single-tree replicates were used per treatment.
respiration, and specific leaf weight (SLW) were lower for leaves Morphological data were determined on September 11 and
grown in the shade than those grown in the sun. The photosyn 12, 1979, and included length of scaffolds, laterals, and inter
thetic efficiency of leaves in the inner shade zone of an apple nodes, node number, stem diameter, and average leaf area of
tree under orchard conditions was determined to be about one- leaves formed after shade treatment was applied, as well as SLW
third that of those in the outer, full-sun zone (9, 15). (of both leaves expanded before and after shade treatment was
Net photosynthetic rates of 9.5 to 15 mg C 0 2 dm -2h r- 1have applied). Chlorophyll was determined by the method described
been reported for peach leaves (6, 7). Crews et al. (7) reported by Arnon (1).
a light compensation point below 1 mW cm -2 (46 |xE s -1 m -2 ) Gross photosynthesis and stomatal resistance were measured
of PAR for sun leaves; no data have been reported for leaves with a ventilated diffusion porometer (Model VP-1, Cayuga
grown in shade. Development, Ithaca, N.Y.) in full sun using the method de
The objective of this study was to determine the effects of scribed by Peet et al. (16). Photosynthesis was measured while
varying light conditions on the growth, morphology, and leaf exposing the abaxial surface of the leaf (1 cm2) to ,4C 0 2 (10.1
photosynthetic characteristics of peach trees. jxCi/liter, 334 ppm C 0 2, 21% 0 2, 19 ml) for 30 sec. Simul
taneous stomatal-resistance determinations were made.
Materials and Methods Immediately after pulsing, the exposed area was excised with
One-year-old peach trees (‘Redhaven’ on Halford rootstock) a No. 11 cork borer and placed in a scintillation vial containing
were pruned to one 20-25 cm shoot and planted in 38-liter pots 0.5 ml of Protosol tissue solubilizer (New England Nuclear) and
in 1 sand : 1 sphagnum peatmoss : 1 soil (by volume) on May digested for at least 48 hr. Samples were bleached with 1.0 ml
5, 1979. Fertilizer, pesticides, and water were added as required. of benzoyl peroxide in toluene (5 g in 30 ml). After 24 hr, 15
Trees were under full sun until June 20, 1979, then placed under ml of scintillation fluid (5 g PPO/liter of toluene) was added
artificial shade treatments that gave 100%, 36%, 21%, and 9% and radioactivity was determined with a Beckman LS 100 Liquid
of full sun. The leaf below the terminal was tagged to distinguish Scintillation spectrometer. Corrections were made for back
pre- from postshade treatment leaves. Full solar radiation was ground and quenching and gross photosynthetic rate was cal
culated— as mg C 0 2 dm -2hr-1 , mg C 0 2 mg chlorophyll- ‘hr-
or mg C 0 2 mg dry w t- ‘hr - 1— by using leaf disc area, exposure
Received for publication April 21, 1982. Michigan Agricultural Experimental
time, radioactivity, specific activity of C 0 2, SLW, and chlo
Station Journal Article No. 10315. The cost of publishing this paper was defrayed
in part by the payment of page charges. Under postal regulations, this paper rophyll values.
therefore must be hereby marked advertisement solely to indicate this fact. Net photosynthetic rates and light-response curves were de
'Present address: Department of Horticultural Science, University of Guelph, termined with an open gas analysis system on whole leaves on
Guelph, Ont., Canada NIG 2W1. July 16, 1980, as previously described (19). Determinations
were made on the first fully expanded leaf after treatments at
25° ± 0.5°C, ambient C 0 2, 0 2 and N2 levels, and a range of
PAR levels (see Fig. 1).
Sunlight Chlorophyll Chlorophyll Total
Results (%) a b Chlorophyll « a/b
The shading treatments did not affect shoot length, internode mg! dm2
length, or node number (Table 1). Stem diameter decreased as 100 2.8 az 1.3 a 4.2 a 2.2
shading increased, but only heavy shade (9% full sun) had a 36 4.9 b 1.7 b 6.6 b 2.8
significant effect (Table 1). Area per leaf was increased 18%, 21 4.8 b 2.2 c 7.0 b 2.2
30%, and 20% for 36%, 21%, and 9% full sun, respectively, 9 5.2 b 2.5 c 7.7 b 2.1
(Table 1), but the difference was significant only at 21% full fig chlorophyll!mg dry wt
sun. As shading increased, we observed that leaves and shoots 100 2.6 a 1.2 a 3.8 a 2.1
became more horizontal. 36 6.1 b 2.2 b 8.3 b 2.8
SLW decreased as PAR decreased for leaves initiated before 21 7.1 b 3.4 c 10.9 b 2.1
9 12.1 c 5.7 d 17.9 c 2.1
(preshade) treatments were applied (Table 1). For 36%, 21%,
and 9% full sun, SLW decrased 22%, 31%, and 47% in preshade 7Mean separation within columns and.parameters by Duncan’s multiple
leaves and 28%, 42%, and 61% in postshade leaves, respec range test, 1% level.
tively.
Chlorophyll content in leaves formed after shade was applied cantly. When the photosynthetic rate of the postshade leaves
increased as light decreased, whether expressed on the basis of was expressed as mg C 0 2 per mg chlorophyll, the rate decreased
surface area or dry weight (Table 2). Chlorophyll a, chlorophyll as shading increased; but when it was expressed on a per mg
b, and total chlorophyll were significantly higher under shade dry-weight basis, it increased.
than under 100% full sun, but the ratio of chlorophyll a to Stomatal resistance was not affected significantly by shading
chlorophyll b was not affected significantly by shading. (Table 3). Light saturation occurred between 700 and 900 jjlE
The photosynthetic rate per unit area of postshade treatment s _1m ~ 2 for 100% full-sun leaves, while 9% full-sun leaves
leaves decreased as shading increased, but the reduction was became saturated between 400 and 600 |xE s _1m ~ 2 (Fig. 1).
significant only at 9% full sun (Table 3). Shading treatments The light compensation point was below 100 (jlE s -1 m “ 2 for
did not affect photosynthetic rate of preshade leaves signifi- both treatments.
Discussion
Response of peach to various levels of solar radiation paral
leled that previously reported for apple: shading decreased stem
diameter, increased leaf area, and decreased specific leaf weight.
Maggs (14) observed that shading of apple leaves reduced length
of new stems, number of leaves, intemode length, and total leaf
area. Area per leaf was greatest in intermediate shade (21% full
sun) and total weight per leaf was greatest in full light. Priestley
(17) found that shade (V6 to !/io daylight) increased specific leaf
area (inverse of SLW) in apple. Jackson and Palmer (13) reported
that shading reduced the number and weight of new apple shoots,
the fresh weight per unit length of shoot, girth increment, and
leaf thickness and weight per unit area. In our work, shading
did not result in as many morphological changes as reported for
apple, but leaves and branches were more horizontal and leaves
were larger, increasing the effective light interception area per
Fig. 1. Photosynthetic light response curves for leaves of ‘Redhaven’ tree.
peach that expanded under 100% or 9% of full sun. These are SLW in apple was reduced by low light (2). We obtained
representative curves and each point is the mean of 2 leaves per similar results for leaves formed both before and after the shade
tree. was applied. Barden (2) hypothesized that SLW might be a useful
Table 1. The effect of shade on scaffold length, intemode length, node number, scaffold diameter, leaf area, and
specific leaf weight of ‘Redhaven’ peach7.
Scaffold Intemode length Scaffold Avg Specific leaf wt
Sunlight length _______ (cm)_______ ___ Node number___ diameter leaf area _____ (mg/cm2)
(%> (cm) Preshade Postshade Preshade Postshade (mm) (cm2/leaf) Preshade Postshade
100 36.8 1.9 1.5 13.8 11.6 4.1 a 29.6 a 10.1 a 11.1 a
36 44.8 1.6 1.7 15.8 11.0 3.9 ab 34.9 ab 7.4 b 8.0 b
21 44.0 1.6 1.8 14.9 11.9 3.6 ab 38.6 b 7.0 b 6.4 be
9 41.0 1.5 1.8 16.1 8.7 3.1 b 35.6 ab 6.4 c 4.3 c
zMean separation within columns by Duncan’s multiple range test, 5% level.
Photosynthetic rate
Preshade Postshade Postshade Stomata resistance
Sunlight (mg C 0 2 (mg C 0 2 (mg C 0 2/mg (lag C 0 2/mg (s c m '1)
(%> dm -2h r~ ') dm “ 2h r_1) chlorophyll h r-1 ) dry wt hr - 1) Preshade Postshade
100 18.8Z 23.2 ay 5.6 ax 2.1 ay 4.9Z 4.6Z
36 22.2 22.7 a 3.4 b 2.8 b 4.4 3.8
21 20.6 19.7 a 2.8 be 3.1 b 4.8 5.5
9 16.5 12.7 b 1.7 c 3.0 b 5.3 7.9
zNo statistically significant differences.
yMean separation within columns by Duncan’s multiple range test, 5% level.
xMean separation within columns by Duncan’s multiple range test, 1% level.
index of previous light exposure and photosynthetic potential. centration of 14C 0 2 throughout the leaf is zero. Also, in some
This appears to hold for our postshade leaves, but not for our tissues, air spaces or grooves may trap large volumes of air, and
preshade leaves. therefore, unassimilated ,4C 0 2.
The chlorophyll content (Table 3) increased with decreasing The previous history of the leaf in terms of light exposure can
light intensity in our studies. Boardman (4) reported that leaves have a profound effect on individual leaf photosynthetic poten
grown under low PAR had more chlorophyll per unit weight or tial. If the total photosynthetic potential of a tree is to be esti
unit volume of leaf, but that chlorophyll content per leaf area mated, we must integrate information on the previous light
is very often lower than that of leaves grown in higher PAR. microclimate with present conditions.
Our results do not totally agree with Boardman’s in that chloro
phyll increased with decreasing light on both an area and dry- Literature Cited
weight basis in peach leaves.
1. Amon, D.L. 1949. Copper enzymes in isolated chloroplasts: po-
As shading increased, chlorophyll content per unit area in lyphenoloxidase in Beta vulgaris. Plant Physiol. 24:1-15.
creased, but photosynthetic rate per unit area decreased. When 2. Barden, J.A. 1974. Net photosynthesis, dark respiration, specific
expressed on a dry-weight basis, both the rate of C 0 2 fixed and leaf weight, and growth of young apple trees as influenced by
chlorophyll content increased as PAR decreased. However, C 0 2 light regime. J. Amer. Soc. Hort. Sci. 99:547-551.
assimilation per unit chlorophyll decreased with shading, sug 3. Bjorkman, O. and P. Holmgren. 1966. Photosynthetic adaptation
gesting that the light reaction (chlorophyll) in shade leaves was to light intensity in plants native to shaded and exposed habitats.
not as efficient as that in sun leaves or that the dark reaction Physiol. Plant. 19:854-889.
was limiting. Leaf morphology, chloroplast structure, mesophyll 4. Boardman, N.K. 1977. Comparative photosynthesis of sun and
resistance, or RuDP carboxylase activity may be limiting pho shade plants. Annu. Rev. Plant Physiol. 28:355-377.
tosynthesis in these light-stressed leaves. 5. Cain, J.C. 1971. Effects of mechanical pruning of apple hedge
rows with a slotting saw on light penetration and fruiting. J. Amer.
The light compensation point for both the 100% full-sun and Soc. Hort. Sci. 96:664-667.
9% full-sun leaves was very low (below 100 |xE s~ 'm -2 ). The 6. Chalmers, D.J., R.L. Canterford, P.H. Jerie, T.R. Jones, and
maximum rate of photosynthesis was much lower for 9% full T.D. Ugalde. 1975. Photosynthesis in relation to growth and
sun than for 100% full sun (Table 3, Fig. 1). The rate of C 0 2 distribution of fruit in peach trees. Austral. J. Plant Physiol.
assimilation of leaves growing on the inside of a peach tree (at 2:635-645.
lower light levels) would be lower than leaves growing at the 7. Crews, C.E., S.L. Williams, and H.M. Vines. 1975. Character
top of the tree, even though the former would be operating at istics of photosynthesis in peach leaves. Planta 126:97-104.
their maximum rate. Crews et al. (7) reported that photosynthetic 8. Heinicke, D.R. 1964. The microclimate of fruit trees: 111. The
rate for peach leaves increased linearly with PAR until saturation effect of tree size on light penetration and leaf area in Red De
licious apples trees. Proc. Amer. Soc. Hort. Sci. 85:33-41.
was reached and Chalmers et al. (6) found different photosyn
9. Heinicke, D.R. 1966. Characteristics of McIntosh and Red De
thetic rates throughout peach tree canopies. However, the pre licious apples as influenced by exposure to sunlight during the
conditioning light levels, which would determine the maximum growing season. Proc. Amer. Soc. Hort. Sci. 89:10-13.
potential photosynthetic rate of each leaf, were not determined. 10. Incoll, L.D. 1977. Field studies of photosynthesis monitoring with
A leaf growing low in the tree would have a lower maximum 14C 0 2, p. 137-155. In: J.J. Landsberg and C.V. Cutting (eds.).
photosynthetic rate than a leaf from the upper, outer edge, even Environmental effects on crop physiology. Academic Press, Lon
if both were light-saturated. don.
The photosynthetic rates determined by the UC 0 2 method 11. Jackson, J.E. 1978. Utilization of light resources by high density
(Table 3) and the net photosynthetic rates (Fig. 1) are different. planting systems. Acta Hort. 65:61-70.
The maximum net photosynthetic rates at 100% full sun (around 12. Jackson, J.E. 1980. Light interception and utilization by orchard
system s. H ort. Rev. 2 :2 0 8 -2 6 7 .
12 mg C 0 2 dm ~2hr -1) are in the range of those found by Crews 13. Jackson, J.E. and J.W. Palmer. 1977. Effect of shade on growth
et al. (7), viz., 9 to 12.4 mg C 0 2 dm -2 h r-1 . Thus, the gross and cropping of apple trees: I. Experimental details and effects
photosynthetic rates may be overestimated. Incoll (10) outlined on vegetative growth. J. Hort. Sci. 52:245-252.
several sources of error, using the 14C 0 2 method, that could 14. Maggs, D.H. 1960. The stability of the growth pattern of young
overestimate the photosynthetic rate. A higher diffusion rate for apple trees under four levels of illumination. Ann. Bot. 24:434-
14C 0 2 exists at the start of an exposure because the initial con 450.
15. Mika, A. and R. Antoszewski. 1972. Effect of leaf position and rootstock varieties under reduced illumination. Ann. Bot. 33:967—
tree shape on the rate of photosynthesis in the apple tree. Pho- 980.
tosynthetica 6:381-386. 18. Proctor, J.T.A ., W.J. Kyle, and J.A. Davies. 1975. The pene
16. Peet, M.M., A. Bravdo, D.H. Wallace, and J.L. Ozbun. 1977. tration of global solar radiation into apple trees. J. Amer. Soc.
Photosynthesis, stomatal resistance, and enzyme activities in re Hort. Sci. 100:40-44.
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J. Amer. Soc. Hort. Sci. 108(4):544-548. 1983.
Crop Productivity Efficiency of Petunias
in the Greenhouse
Richard H. Merritt1 and Harry C. Kohl, Jr . 2
Department of Environmental Horticulture, University of California, Davis, CA 95616
Additional index words, photosynthetic efficiency, Petunia hybrida, leaf area index, source-sink relationships, root zone
temperature
Abstract. Crop productivity efficiencies (CPE) of around 8% (the ratio of the dry weight gain of the crop to the
potential to produce dry weight), were realized with petunias (Petunia hybrida Villm.), provided that the crop canopy
was essentially closed at the beginning of the 9- to 12-day experimental periods and that there were many branches
(sinks). This was found at either long or short photoperiods or at either a normal (15.6°C) or reduced (7.2°) temperature
for the 16-hour night periods. Long photoperiods resulted in significantly increased CPE through increased size of
the leaves before the crop canopy was closed. Elevated root temperature increased CPE after a sizeable number of
lateral branches had formed.
Greenhouses require high plant production per unit area in during the day photosynthetic photon flux, carbon dioxide ad
order to be profitable. Fundamental to high plant production is equacy, and photosynthetic enzyme adequacy were the same for
high dry weight gain per unit area. The authors suggest that both crops.
there are 7 primary crop parameters which determine dry weight In another study, a petunia crop grown at a long photoperiod
gain at a given light flux. Present and proposed crop investi showed increased dry weight production in comparison with a
gations should consider these parameters, which are as follows: short-photoperiod crop only when the light-gathering capacity
a) The effectiveness of the crop in absorbing the incident light of the crop was below the maximum and before a large number
is the leaf canopy efficiency (LCE). The most interesting and of sinks were available, i.e., before branching had occurred (5).
manageable aspect of this is the leaf area index which is the Increased root temperature caused increased dry weight produc
area of leaves per unit area of greenhouse bench (LAI). tion only after the young plants had branched and thereby greatly
b) Sinks adequacy (SKA) for accepting photosynthates which increased the number of sinks per unit area.
is usually highly correlated with the number and activity of shoot This paper reports the results of a study of leaf canopy effi
apices per unit of cropped area. ciency and sink adequacy using young petunias as the test crop.
c) The adequacy of C 0 2 supply at the chloroplast level which The environmental conditions varied were photoperiod, night
includes consideration of photorespiration. temperature, and root temperature. To account for small differ
d) Photosynthetic enzyme adequacy. ences in photosynthetic photon flux (PFF) which the crops re
e) Synthesis respiration costs which when measured as a loss ceived and to allow for a wider comparison of results, a crop
in dry weight are high for fats and proteins, low for starches productivity efficiency (CPE) was calculated. CPE is the ratio
and cellulose, and negative for organic acids. of the dry weight gain of the crop to the potential to produce
f) Maintenance respiration cost which is a direct function of dry weight. The potential to produce dry weight in grams was
the growing biomass of the crop. calculated by dividing PPF in kilojoules by 14.6 kilojoules which
g) Depletion cost which accounts for dry weight loss by such is the heat of combustion of 1 gram of glucose. PPF in kilojoules
phenomena as harvest, predation, and abscission. was obtained by multiplying PPF in moles by 217.4 (1). The
In previous studies it was reported (2, 3) that the dry-weight results are explained by larger leaves at the long photoperiod
gain o f a chrysanthem um crop grow n at a low night tem perature providing both more nearly adequate sinks and a more rapid
was at least equal to that of a crop grown at normal night tem increase in the capacity to intercept light.
perature provided that: (a) the light-gathering capacity was high; Throughout the paper there is continual reference to “ petunia
(b) there were adequate sinks (200 stem apices per m2); and (c) crop(s)” rather than “ petunias.” This is done purposefully to
focus the reader on the crop community rather than the plant.
Received for publication July 1, 1982. The cost of publishing this paper was
defrayed in part by the payment of page charges. Under postal regulations, this
paper therefore must be hereby marked advertisement solely to indicate this fact. Materials and Methods
'Visiting Professor from Department of Horticulture and Forestry, Cook College,
Rutgers University, New Brunswick, NJ 08903. Young, healthy, well-branched, greenhouse-grown seedlings
2Plant Physiologist, California Agr. Expt. Sta., Davis, CA 95616. of ‘Snow Cloud’ petunia in 7.6-cm, square plastic pots were
