plantas con estrés por sequía.
Ambos controles y las
Efecto del estrés por
sequía en las
características
fotosintéticas, el
crecimiento y la
acumulación de
azúcares del sorgo
dulce cultivado en el
campo.
ANGELO MASSACCI, ALBERTO BATTISTELLI AND FRANCESCO
LORETO
Resumen
Estudiamos el efecto de la sequía en la fotosíntesis, el
crecimiento y la acumulación de azúcar en el sorgo
dulce cultivado en el campo. Los estomas se cerraron
cuando el estrés por sequía se desarrolló durante el día.
Como consecuencia, la fotosíntesis se redujo. La
eficiencia en el uso del agua de las hojas estresadas por
la sequía fue ligeramente menor que la de los controles
y esta reducción fue más pronunciada temprano en la
mañana y en la noche. Esto sugirió que no solo los
estomas limitan la fotosíntesis bajo la sequía: la
fotosíntesis a altas concentraciones de CO2 fue más
baja en la sequía que en las hojas testigo, lo que sugiere
que el metabolismo del carbono se inhibió. El
transporte de electrones también se redujo, pero
pareció ser necesario un mayor número de electrones
para fijar el CO2 en condiciones de sequía. Antes de la
antesis, el crecimiento no se vio afectado por la sequía,
pero después de la antesis, la elongación de los últimos
entrenudos fue mayor en los controles que en las
plantas estresadas por la sequía acumularon sacarosa y
almidón en el tallo después de la antesis. La
acumulación fue baja y cesó temprano en el ápice del
tallo. Es probable que las semillas en la panícula
compitan con el ápice de carbono. Durante la
maduración de la panícula, el contenido de sacarosa y
almidón fue mayor en sequías estresadas que en tallos
de control. Al final de nuestros experimentos, sin
embargo, el contenido de sacarosa por planta fue
similar en todas las muestras. Concluimos que la
acumulación de azúcar es menos sensible a la sequía
que la fotosíntesis. Independientemente de las
condiciones de sequía, (i) antes del crecimiento de la
antesis es el sumidero predominante para los
fotosintéticos; (ii) después de la acumulación de
azúcares antesis en convertirse en el sumidero
preferencial a expensas del crecimiento de los
entrenudos apicales.
Introducción
Sorghum is a drought-resistant C4 plant and is
cultivated mainly during the dry season in warm areas
of the world. Drought resistance has be attributed to
high stomatal sensitivity to drought stress (Ackerson et
al. 1980). Stomatal closure occurs when soil water
content is low or in response to high evaporative
demand. Stomatal sensitivity, however, is gradually lost
after flowering (Ackerson et al. 1980; Garrity et al.
1984). As a consequence, the water use efficiency (i.e.
the ratio between leaf photosynthesis and
transpiration) is reduces and drought stress may
negatively affect again filling and development
(Premachandra et al. 1994). A positive relationship has
been found between leaf photosynthesis, total biomass
and grain production in the presence or in absence of
water limitation (Peng et al. 1991). Therefore, irrigation
greatly improves the yield of grain sorghum cultivars,
particularly if dry conditions occur after flowering.
The physiology of sweet sorghum is different from that
of grain and fibre sorghum. Sweet sorghum cultivars are
utilised to produce sugar, ethanol, syrups and molasses.
The sugar, mainly sucrose, is accumulated in large
amounts in the stem during the development of the
inflorescence (McBee ad Miller 1982). During this
period, there is no competition between gain
development and sugar accumulation (Lingle 1987).
Sucrose accumulation in the stem is associated with a
decreased activity of sugar-degrading enzymes, on the
other hand, is also associated with reduction of stem
elongation in maize (Setter and Meller 1984) and sweet
sorghum (Lingle 1987).
We have investigates the effect of drought stress on
thee physiology of field-grow sweet sorghum with
particular emphasis on the accumulation of sucrose and
on the effects on growth. Although stress was imposed
over the whole growth period, we focused our
observation on the period of grain development
because of the coincidence between panicle
maturation, maximum sensitivity to drought stress, and
sugar accumulation in the stem.
We chose as a case-study sweet sorghum cv. Korall,
because previous experiments on this cultivar indicated
a low sensitivity of photosynthesis to drought stress
(our experimental results and J. Woods, unpublished).
Our experiments confirmed that sweet sorghum cv
Korall is less sensitive to drought stress than a cultivar
of fibre sorghum (H-29) characterised by a similar
vegetative cycle and cultivated under the same
environmental conditions. Drought stress negatively
affected the photosynthesis and stomatal conductance
of Korall leaves but photosynthesis was still appreciable
at very low leaf water potentials. In drought- stressed
plants, growth was reduced only after flowering and
sugar content per plat was almost unchanged relative to
what was found in irrigated plants. We conclude that,
contrary to what is found in grain and fibre sorghum,
drought stress may affect plan carbon metabolism but
seems to have a minor impact on sweet sorghum
productivity.
Material and Methods
Plat Material and Experimental Conditions
The experiments were carried out at the CNR-IBEV
experimental field of Moterotondo, near Rome, Italy,
during summer 1994. The summer was dry and no rain
fell except during the first month of growth (June).
Seeds of sweet sorghum cv. Korall and of fibre sorghum
cv. H-29 were planted in loamy, fertilised soil. Each
experimental condition was repeated in four plots. Each
plot was 6 x 4 m; seed spacing was 0.7 m between rows
and 0.15 m in the row. Plots were distributed according
to a random experimental design. Potential
evapotranspiration was calculated from
micrometeorological data using a Penmann equation
modified for sorghum stands. Real evapotranspiration
was measured with a class A pan evaporimeter. In the
plots of control plants, real evapotranspiration was fully
compensated by sprinkle irrigation. Plants were
generally irrigated every other day during the final
growth stages in order to maintain a leaf water
potential (Ѱ) higher that -1.5 Mpa along the day. In
plots of drought-stressed plants, water deficit was not
compensated by irrigation plants were rainfed only
during the first month of growth. Fifty plants were also
grown in commercial soli in 50-L pots (one plant per
pot). Some of these plants were regularly irrigates
during the experiment, other were gradually stressed by
withholding water after anthesis.
As a stress index we used the midday rather than the
pre-dawn leaf water potential because stomata are
insensitive to water potential as high as those measured
at pre-dawnn (Osmond et al. 1980). Moreover, the
midday water potential is considered an useful indicator
of the conditions of the soil- plant-atmosphere
continuum in sweet sorghum (G. Gosse, personal
communication) since it integrates the daily evaporative
demand and the soil water status (Cary and Wright
1971). Leaf water potential was measured on fully
expanded leaves with a pressure chamber (PMS,
Corvallis, Oregon). Leaf water potential was also
measured while measuring the diurnal trend of
photosynthetic parameters in control and drought-
stressed leaves.
Measurements of Photosynthetic Parameters
Photosythesis ad stomatal conductance were measured
in the field with a LiCor 6200 (LiCor, Lincoln, Nebraska)
portable gas analyser o samples randomly selected in all
plots. The photosynthetic water use efficiency (WUE)
was calculated by dividing leaf photosynthesis by the
transpiration rate. Measurements were carried out at
several growth stages on the last fully expanded leaf.
However, only measurements carried out on the flag
leaf beaten anthesis and pinnacle maturation are
presented with the exception of Fig. 3. During the
diurnal trends each measurement was replicated o five
randomly chosen leaves exposed to the same
photosynthetic photo flux density (PPFD) ad with similar
leaf temperature. The order of leaf sampling was also
randomised.
Measurements of photynthesis ad stomatal
conductance were also made on potted plants using a
laboratory gas-exchange system described by Loreto et
al.(1992). This system allowed measurement of the
response of photosynthesis to different irradiances ant
to calculate the quantum yield of CO2 fixation (фco2). It
also allowed measurement of response of
photosynthesis to the intercellular CO2 partial pressure
(ci). This parameter was measured also I the field. In this
case, the CO2 partial pressure was temporarily enriched
by breathing into the cuvette (Mc Dermitt et al. 1989).
The equations of von Caemmerer and Farquhar (1981)
were used for calculating the photosynthetic
parameters.
In the laboratory, chlorophyll flourescence emission and
gas exchange could be measured simultaneously as
already described (Loreto et al. 1992) using a PAM-101
(Walz, Efferlrich, Germay) modulated fluorometer.
Flourescence was also measured I the field, using a
PAM-200 (Walz) portable flourometer. In the field, the
potential yield of photochemistry (Fv/Fm, where Fv is the
variable fluorescence and Fm is the maximum
florescence) was estimated by dark adapting the leaves
for 20 mi. In the laboratory, the quantum yield of the
electron transport through PSII (фPSII) was estimated
using the equation of Genty et al. (1989):
ФPSII= F/ F’ m ,
Where F is the difference between the maximum
fluorescence observed under actinic light (F’ m) and the
fluorescence measured under steady state conditions
(Fs). Other details of fluorescence parameters and
measurements are reported in Loreto et al (1992).
Growth Analysis
On-destructive analysis of plants growth was carried out
from emergence to the grain black layer stage on Korall
plats irrigated or drought-stressed. Measurements of
plant total leaf area, leaf number, stem height and
width are reported. Each data point is the average of at
least 10 measurements on different plants sampled
randomly in the different plots.
Measurement of Sugar Content
Five control and five drought-stressed plats were
randomly harvested I the experimental plots. Harvest
was made in four different periods between anthesis as
the black layer stage. A piece of stem 5 cm long was cut
at three different heights: 10 cm from soli (basal stem),
100 cm from soil (central stem), and 10 cm from the
panicle (apical stem). The external layer or green
parenchyma was plainly removed in order to avoid
interference caused by pigments during sugar analysis
and the samples were frozen in liquid nitrogen and
maintained at -80°C until extracted. Before extraction,
50 g subsamples were ground in a glass-glass
homogenizer. Extraction was doe maintaining the
samples for 45 min in 2 mL of 80% ethanol, 20% water
buffer containing 100 mM Hepes-KOH (pH 7.1) and 1o
nM MgCl2 al 80°C. The extract was cooled at room
temperature and centrifuged at 15800 g for 5 min.
Soluble sugars I the supernatant were analyzed
enzymatically according to jones et al. (1997) using an
anthos 2001 plate reader (Anthos Labtec Istrum.,
Salzburg, Austria) at 340-405 nm. The pellet was re-
suspended and washed at least four times with 40 mM
acetate buffer (pH 4.5) and the autoclaved at 120°C for
40 min to solubilize starch. To complete starch
hydrolysis, 4 U of α-amylase and 40 U of
amyloglucosidase were added ad the mixture was
incubated for 1 h at 50°C. After starch hydrolysis,
samples were centrifuged and glucose in the
supernatant was analysed as previously done for
soluble sugars. The contents of sucrose ad starch are
reported as glucose equivalents. Recovery of added
carbohydrates was higher than 90%. Differences
between means ad interactions between treatments,
sampling position and sampling period on sugar content
were tested by ANOVA.
Results
Effect of drought stress on photosynthesis
(a) Water potential
Photosynthesis was progressively reduced at decreasing
leaf water potential (Fig. 1). The best-fit lines were
significantly different I the cultivars and indicates that
photosynthesis of Korall leaves is generally less affected
by growing water tress than photosynthesis of H-29
leaves. Eve when Ѱ was lower than -2.0 MPa, Korall
leaves were still photosynthesising at a rate of about 18
μmol m-2s-1. On the other hand, photosynthesis of H-29
leaves was totally inhibited in leaves with a Ѱ around -
2.0 MPa. The response to drought stress of field-grow
plants was not different from that of potted plants both
in cv. Korall and H-29.

(b) Diurnal trend
Diurnal trends of photosynthesis and stomatal
conductance were measured once a week between
anthesis and the black layer stage. Since the daily
changes were similar, only the diurnal trends measured
o flag leaves at the beginning of anthesis and at panicle
maturation are presented. These data were collected
simultaneously with the first and the last sugar
sampling, respectively.
Fig.1. Response of photosynthesis to decreasing leaf water
potential I sweet sorghum cv. Korall, (open symbols) and in
fibre sorghum cv. H-29 (closed symbols). Measurements were
carried out in field-grown (circles) and potted (squares) plants
at midday under saturating PPFD and ambient CO2 partial
pressure. Second-order best-fit lines show the different
trends of reduction of photosynthesis at increasing leaf water
potential in the two cultivars. Confidence limits (95%) of the
lines were significantly different.
Maximum photosynthesis was reached during the
middle hours of the day both in drought-stressed and
control leaves but photosynthesis of drought-stressed
leaves was on average 65% that of control leaves (Fig.
2a). This difference was even more evident at the
beginning of the following day: gs was very sensitive (Fig
2c). During the day, gs of drought-stressed leaves was
40% that of control leaves, but the reduction was less
evident in the evening and in the early morning, when
gs of control leaves was also low. Midday depression of
photosynthesis, associated with temporary stomatal
closure during the warmest and less humid hours of the
day, was never found in control and drought-stressed
Korall leaves. The photosynthetic WUE was higher in
control leaves than in drought-stressed leaves. This
difference was particularly evident at the beginning and
at the end of the day. During the day, changes of WUE
occurred in drought-stressed leaves during the central
hours of the day (Fig. 2c). The gs measured at different
growth stages was related to an index which includes
photosynthesis, atmospheric pressure, co2 partial
pressure, and relative humidity (Fig. 3). The slope of this
relationship did not change during the season and was
not different I control and drought-stressed plants.
Fig. 2. Diurnal trend of photosynthesis (a), stomatal conductance (b), photosynthetic water use efficiency
(photosynthesis/transpiration) (c) and leaf water potential (d) of control (open symbols) and drought-stressed (closed symbols)
leaves. Measurements shown were carried out at the beginning of anthesis (triangles) and during panicle maturation (circle) in field-
grow plants. Each data point is an average of five measurements taken at the same PPFD. Bars = s.e. When not visible, s.e. is smaller
than symbol size. PPFD and leaf temperature changed similarly during the two diurnal trends shown.

(c.) CO” response photosynthesis of drought-stressed leaves satured at

much lower ci than photosynthesis of control leaves (Fi.
Photosynthesis of control and drought-stressed leaves
4b)
was similar at very low ci (Fig. 4). Field measurements
frequently indicated a very strong reduction of ci (d) Photochemistry and electron transport
because of stomatal closure. Under this condition,
The photochemical efficiency was temporarily reduced
saturation of photosynthesis was not attained (Fig. 4a).
by drought stress. At midday in the field, the
When we measured the photosynthetic para meters
photochemical efficiency, as determined by the
with the laboratory equipment, we able to increase the
fluorescence ratio Fv/Fm, was lower in drought-stressed
CO2 partial pressure in the cuvette where the leaf was
than in control leaves. However, in the early morning
enclosed up to 2000 μbar. It was then clear that
(at Ѱ always higer tha -1.2 MPa), the photochemical
efficiency was about 0.75 both I control and, drought-
stressed leaves (not shown). The quantum yield of CO2
fixation (i.e. mol. of CO2 fixed per mol. of quanta of
incident light during the day) was also reduced in
drought stress conditions (see also Fig. 1). To see how
the relationship between photochemical reaction ad
carbon metabolism was affected by drought stress, the
ratio between фPSII and фCO2 was measured at a PPFD
comparable to that experienced in the field I leaves
characterized by different water potentials (Fig. 5). The
slow rise of this ratio indicates that drought stress
affected фCO2 and фPSII to different degrees.
Fig. 3. Relationship between stomatal conductance (gs) and
the index formed by relative humidity at the leaf surface (rh),
photosynthesis (A), atmospheric pressure (P) and ambient
partial pressure of CO2 (ca). Measurements were carried out
during vegetative growth (●) ad between anthesis and panicle
maturation (○) o control and drought-stressed plants.
Effect of Drought Stress on Growth Rate
The final number of leaves set was slightly higher in
control plants than in drought-stressed plants (Fig. 6a).
A similar trend was observed for the total leaf area (Fig.
6b). Stem length was affected by drought stress only
during the last month (Fig 6c). During panicle
maturation, control plants extended last internodes
significantly more than stems of drought-stressed
plants. Stems were slightly wider in drought-stressed
plants than in control plants throughout the period of
vegetative growth (Fig. 6d). However, stem width of
control and drought-stressed plants was similar at
panicle maturation.
Effect of Drought Stress on Stem sugar Content and
Accumulation
Sucrose, starch and hexose contents were measured
four times between anthesis and panicle maturation in
the stems of drought-stressed and control plants (Fig.
7). The water content in the stem was similar in
drought-stressed and control plants (76 and 78% of the
total weight, respectively). Therefore, we expressed the
sugar content on a fresh weight basis as it was
measured with the biochemical assay. Sucrose and
starch contents expressed as glucose equivalents were
higher inn drought-stressed tha in control stems. The
difference in sucrose and starch content between
drought-stressed ad control leaves was significant at
P<.005, except at the last sampling time when no
difference was found (Fig. 7a, b). Both sucrose and
starch content increased during panicle maturation and
the increase was particularly evident between anthesis
and grain filling (first and second sampling period). In
drought-stressed stems, sucrose and starch reached
their highest content earlier than in control stems.
Glucose and fructose contest were much lower than
sucrose content and declined during panicle
maturation. The reduction was particularly evident
between the second and third sampling period. Hexose
contents were not significantly different in drought-
stressed and control sample (Fig. 7c, d).
Sucrose and starch were also measured in the basal,
central and apical part of the stem during panicle
maturation (Fig. 8). There was no significant difference
between the sugar content of drought-stressed and
control stems sampled at the same stem height.
Therefore, data for drought-stressed and control
samples were pooled in Fig.8. The highest sucrose
content was found in the central part of the stem but
the difference with the content found in the basal part
was small and not significant. Sucrose accumulation in
the central and basal stem increased during the whole
sampling period. The content of sucrose in the apical
part of the stem was lower than in the other two parts.
In addition, sucrose accumulation in the apical part
ceased between the second and third sampling period,
i.e. during grain filling (Fig. 8a). The accumulation of
starch followed a pattern similar to that of sucrose.
However, the total amount of starch was about 10
times lower than that of sucrose and there were no
differences between the positions of accumulation
during the first two sampling periods (Fig. 8b).
 Fig. 5. Changes in the ratio between the quantum yield of PSII
electron flow (фPSII) and the quantum yield of CO2 fixation
calculated per incident PPFD (фCO2) at decreasing water
potential. All measurements were done after anthesis on
potted in the laboratory under controlled conditions (leaf
temperature 25°C, vapor pressure difference between leaf
and air < 2.0 kPa, ambient CO2 partial pressure)

Discussion
Effect of Drought Stress on Photosynthesis

The photosynthetic apparatus of Korall leaves was

affected by drought stress and the effect was no
relieved overnight.

However,

(1) The photosynthetic rate of Korall leaves was always

(2) Control and stressed leaves did not shown ay
(3) Contrary to what was previously reported on grain
Fig. 4. Response of photosynthesis to the intercellular CO2
partial pressure (ci). Open symbols represent control leaves,
closed symbols are drought-stressed leaves. Each symbol
represents a different leaf. (a) Field measurements. (b)
Laboratory measurements. Leaf water potentials of control
ad drought-stressed leaves were -1.2 ±0.2 and -1.9±0.2MPa,
respectively.
higher than of H-29 at comparable water potential
(Fig. 1); half of the maximum photosynthesis was
maintained in Korall leaves eve when a Ѱ lower
than -2.0 MPa was reached;
midday depression of photosynthesis (Fig. 2). A
depression of photosynthesis concomitant with low
humidity, warm temperature and bright light was
expected when considering the high photosynthesis
of sorghum leaves and was frequently found in
other cultivars during previous experiments;
sorghum (Garrity et al. 1984), stomata did not show
decreasing sensitivity to drought stress during plant
development. In fact, no difference was found in
the response of gs to drought stress during
vegetative growth and panicle maturation. To make
this more clear we used the model reported by Ball
et al. (1987). The slope of the response of gs to the
index obtained considering photosynthesis, relative
humidity, CO2 partial pressure and temperature
represent the sensitivity of gs to these parameters.
Since the slope was the same I measurements
carried out at different growth stage and at
different water potential (Fig. 3), we conclude that
the stomatal sensitivity of Korall leaves did not
change during the season or because of drought
stress conditions.
Stomatal closure effectively maintained a high WUE
during most of the day both in control and drought-
stressed leaves. This contributed to the
maintenance of a favorable water content in plants
during the
 Fig. 6. Measurements of growth of Korall plats in control (open symbols) and drought-stressed (closed symbols) conditions. Each
data point is an average of 10 measurements. Bars = s.e. When not visible, s.e. is smaller tha symbol size. Anthesis occurred 60
days after emergence as shown by the tick in the x-axes.

Hottest hours and allowed photosynthesis to
continue at a moderate rate in drought-stressed
leaves. However, the differences between WUE of
control and drought-stressed leaves at the
beginning and at the end of the day were very clear
and may indicate that stomata were not the only
limitation to photosynthesis in drought- stressed
plants.
Under very stressful conditions (ф<-2.0MPa), stomatal
closure affectively decreased ci to very low values. Field
measurements show that, in some samples, ci may be
so low as to effectively limit photosynthesis (Fig.4a). In
the laboratory experiments, however, we overcame this
limitation by supplying enough CO2 to reach saturation
of photosynthesis (Fig. 4b). It was then evident that
photosynthesis of drought-stressed leaves saturates at
much lower CO2 than photosynthesis of control leaves.
However, photosynthesis of controls and drought-
stressed leaves was similar at low intercellular CO2
partial pressure. This suggests that drought does not
affect the CO2-concetrating mechanism characteristic of
C4 plants. When light is not limiting and there is enough
CO2 to saturate photosynthesis, the photosynthetic rate
of C4 plants is equal to the Vmax of rubisco (Collatz et
al. 1992). The observed response of photosynthesis to
CO2 therefore indicates that a relevant limitation to
photosynthesis of Korall leaves under drought stress
conditions may be caused by a reduced capacity of
rubisco for CO2 fixation. This limitation is probably clear
only when stomatal limitation of photosynthesis is low,
for instance at 7.00 and 18.00 h during the diurnal
trends shown in Fig. 2.
Fig. 7. Contet of sucrose(a), starch(b), glucose (c) and fructose
(d) in the stem of control (open symbols) and drought-
stressed (closed symbols) plants of Korall. Each data point is
the average of 12 samples (4 samples x 3 stem heights: basal,
central and apical). Bars = s.e. When not visible, the s.e. is
smaller (day 0 in the x-axis) to panicle maturation. Sucrose
and starch are expressed in μmol of glucose equivalent for
improving comparison among measurements.
measurement are on a fresh weight (f.w.) basis.
Fig. 8. Sucrose (a) and starch (b) accumulation in the stem of
Korall plats between anthesis (day = in the x-axis) and panicle
maturation. Symbols indicate different position of sampling in
the stem: ●, apical stem, 10 cm lower than panicle. ▪, Central
stem, 100 cm from soil. , basal stem, 10 cm from soil. Each
data point is an average of eight samples (four controls and
four drought-stressed). Bars = s.e. When not visible, s.e. is
smaller than symbol size. Sucrose ad starch are expressed as
glucose equivalent on a fresh weight (f.w.) basis.
We calculated the ratio between the quantum yields of
PSII, estimated from fluorescence measurements
according to Genty et al. (1989), and the quantum yield
of CO2 fixation in field conditions. This ratio estimates
the relationship between electron transport rate and
carbon fixation (Genty et al. 1989; Oberhuber and
Edwards 1993). In C4 plants (and in C3 plats when
photorespiration is suppressed), at least 4 photos are
consumed per CO2 fixed in each one of the two
photosystems; thus the ratio should yield values higher
than 8. In fact, the high demand for ATP on C4 plants
All
further increases the ratio to values higher than 10
(Krall and Edwards 1990). We found that the ratio was
higher than 10 in the absence of drought stress, and
linearly increased with increasing drought stress. This
may indicate that carbon metabolism reactions are
more affected than are photochemical reactions by
drought stress. Carbon metabolism generally down-
regulates the electron transport rate very effectively
(Sharkey et al. 1988); in our experiments, however, this
down-regulation does not seem to be complete and the
progressive changes of the ratio suggests that more and
more electrons are required to fix CO2 when drought
stress is high. It is also possible that the efficiency of
light use for carbon fixation stays unchanged while the
excess of electrons available under drought stress
conditions drives reactions which scavenge oxygen
radicals (Massacci et al. 1995)
Effect of Drought Stress on Growth Rate
Leaf and stem growth of drought-stressed plats were
only marginally reduced with respect to control plats
during the first 60 days of growth (i.e. during vegetative
growth). However, the photosynthetic rate was clearly
reduced during periods of drought stress. We did not
measure root growth and expansion, and are therefore
not able to demonstrate that changes in photosynthesis
are related to changes in carbon partitioning between
shoots and roots. However, this is likely to be the case
because a good correlation between photosynthesis
and total biomass production has been reported in
sorghum plants (Peng et al. 1991). Peng and Krieg
(1992) also showed that a positive correlation exists
between photosynthesis and shoot biomass od leaf area
of grain sorghum. According to our results, however,
this relationship does not exist in sweet sorghum during
vegetative growth. Since no accumulation of sugar in
the stem is evident between germination and anthesis
(Tarpley et al. 1994 and data not shown), it is likely that
vegetative growth is the strongest sink for
photosynthates during the growth period.
Interestingly, after anthesis the elongation of the last
internodes was greater in controls that in drought-
stressed plats. Reduction of internodes elongation was
attributed to a diminished activity of sucrose-degrading
enzymes and, consequently, to an increase of sucrose in
the stem (Setter and Meller 1984; Lingle 1987).
Accordingly, we found a slightly higher content of
sucrose in drought-stressed stems that in control stems.
After anthesis, a reduction of leaf area also occurred
which was caused by falling senescing leaves. In terms
of source-sink interaction, this may represent a
decrease in the availability of photosynthates and
ultimately a reduction of sugar synthesis. The leaf area
reduction occurred few days earlier in drought-stressed
leaves than in controls and may explain why sugar
accumulation stopped earlier in the stem of drought-
stressed plants than in the stem of control plats (Fig. 7).
Effect of Drought Stress on Sugar Accumulation
Given the reduction of photosynthesis, we expected to
find changes in photosynthate partitioning and sugar
content of drought-stressed plants, as also suggested by
Vietor and miller (1990). However, at the end of the
experiment, growth was moderately affected and sugar
accumulation was not affected at all by drought stress
(Fig. 7).
Sucrose accumulates I the stem after anthesis (McBee
and Miller 1982); therefore, we examined the
accumulation of sucrose and starch in different art of
control and drought-stressed stems after anthesis and
during panicle maturation. Plats accumulated 10 times
more sucrose than starch in the stem. The amount of
hexose was comparable to that of sucrose at anthesis.
During panicle maturation, however, accumulation of
sucrose was accompanied by a decrease of hexose
contents in the stems, supporting the idea that sucrose
accumulation is caused by a reduction of sucrose-
degrading enzymes and, consequently, of hexose.
However, the reduction of hexose occurs only after the
second sampling period while most of the sucrose
accumulation occurs between the first and the second
sampling period.
We statistically separated the effect of the three
variables: position, sampling period, and treatment o
sugar accumulation in the stem, and found that the
interaction between the three variable had no
significant effect. However, the interaction sampling
period Х treatment and sampling period Х position were
significant at P<0.005 and P<0.001, respectively. The
first interaction indicates that drought stress caused
sugar to accumulate earlier and, possibly, suggests that
drought stress shortens the vegetative cycle and causes
early senescence of sweet sorghum plants. As a
consequence, sucrose and starch content was higher in
drought-stressed than in control stems at almost all
sampling dates. We followed growth up to the black
layer stage,which has been shown to be the moment
when sucrose content in stem is the highest (Tarpley et
al. 1994). At the black layer stage, the content of We thank Professor R. Leegood and Dr G. Di Marco for
sucrose on a fresh weight basis was not significantly crtical reading of the manuscript. Dr M.A. Iannelli, Dr F.
higher in controls than in drought-stressed stems. Pietrini, Mr D. Tricoli and Mr G. Santarelli provided
However, the content of sucrose and starch was assistance during field and laboratory measurements.
constant only in drought-stressed plats. In control Dr S. Moscatello provided assistance with sugar
plants, sugar content increased until the last sampling analysis. Research supported by Project RAISA, sub-
date. Therefore, we cannot exclude the possibility that project no. 2, paper o, 2512.
sugar continue to accumulate in irrigated sweet
sorghum plants after the black layer stage.
The second interaction indicates that, both I control and
drought-stressed plants, sucrose and starch
accumulated particularly in the central and basal part of
the stem while the lowest accumulation was found in
the apical part of the stem. According to a previous
estimation based on 14C-assimilate partitioning (Vietor
and Miller 1990), the higher accumulation of sucrose is
the lower part of the stem. Our findings localize more
precisely, where the highest sucrose accumulation
occurs. If sucrose distribution in the stem is related to
differences in sucrose-degrading enzymes, as recently
suggested (Tarpley et al. 1994), a similar distribution of
the enzymes may be expected. Interestingly, sucrose
and starch accumulation stops earlier in the apical part
than in the central and basal parts of the stem. These
findings seem to indicate that grain filling is an active
sink of photosynthates, strong enough to complete with
the nearest part of the stem (the apex) for carbon
compounds.
Finally, our results suggest that drought stress may
slightly increase sugar accumulation on a fresh weight
basis, at least until the black layer stage. Using the
growth data we calculated the total amount of sucrose
per plat. We found that control plants accumulated
about 26 g of sucrose per plat stem while drought-
stressed plants accumulated 25 g of sucrose per plant
stem. It is therefore likely that the drought stress
conditions imposed during the experiment do not
perturb sugar synthesis and that, eve under drought
stress condition, sucrose allocation in the stem is the
preferential sink for photosynthates when vegetative
growth has ended and anthesis has occurred.
Since photosynthesis of sweet sorghum plats was less
affected than that of fibre sorghum by drought stress,
we speculate that sugar synthesis is involved in a
unknown mechanism of resistance to the stress.

Acknowledgments