Postharvest Biology and Technology 20 (2000) 63 – 70

www.elsevier.com/locate/postharvbio

Effects of temperature and stem length on changes in

carbohydrate content in summer-grown cut chrysanthemums
during development and senescence
Megumi Adachi *, Saneyuki Kawabata, Ryozo Sakiyama
Laboratory of Horticultural Science, Graduate School of Agricultural Life Sciences, Uni!ersity of Tokyo, 1 -1 Yayoi, Bunkyo-ku,
Tokyo, 113 -8657, Japan

Received 28 June 1999; accepted 17 April 2000

Abstract

Changes in the carbohydrate content in capitula, stems, and leaves were investigated in cut chrysanthemum
Dendranthema×grandiflorum (Ramat.) Kitamura ‘Seiun’ plants with 60-cm stems held at 20, 25 and 30°C for 27 days
postharvest. In addition, plants with 20-cm stems were studied at 25°C. Diameter of the capitula, the angle of the
florets, fresh weight (FW) and dry weight (DW) increased more slowly at 30°C/60 cm and 25°C/20 cm than at
20°C/60 cm and 25°C/60 cm. The number of well-developed florets was notably smaller, and the number of
tubular-shaped florets was larger at 30°C/60 cm than at 20°C/60 cm. Glucose and fructose concentrations in the
capitula increased, although they decreased midway through treatment at 30°C/60 cm. Fructose concentrations in the
stems, and in particular in the middle stems, were greater at 30°C/60 cm than at 20°C/60 cm and 25°C/60 cm. The
increase in the total DW during treatments suggested that photosynthesis was occurring in the leaves and that
photosynthates were a main source for the capitula and stems. Development of the capitula was suppressed, which
was related to the reduction in its carbohydrate supply. Wilting of petals was not accelerated at high temperatures and
with short-stem treatments. © 2000 Elsevier Science B.V. All rights reserved.

Keywords: Cut chrysanthemum; Flower opening; Carbohydrate content; Senescence; Temperature; Stem length

1. Introduction total floricultural production (Funakoshi, 1984).

Chrysanthemums are the most popular cut
flower in Japan and account for one-third of the
* Corresponding author. Present address: People-Plant Re-
lationship Laboratory, Department of Agriculture, Tokyo
University of Agriculture, 1737 Funako, Atsugi City, Kana-
gawa, 228-0011, Japan. Tel./fax: + 81-46-2706541.
E-mail address: ameg@nodai.ac.jp (M. Adachi).
Although cut chrysanthemums have a long vase
life in general, they deteriorate sooner and have a
shorter vase life under high temperatures (Fu-
nakoshi, 1984; Whealy et al., 1987; Cockshull and
Kofranek, 1994; Adachi et al., 1999).
The main cultivar, ‘Shuho-no-chikara’, is usu-
ally produced year-round, except in summer. This
cultivar is sensitive to high temperatures, which

0925-5214/00/$ - see front matter © 2000 Elsevier Science B.V. All rights reserved.
PII: S 0 9 2 5 - 5 2 1 4 ( 0 0 ) 0 0 1 0 6 - X
64 M. Adachi et al. / Posthar!est Biology and Technology 20 (2000) 63–70
accelerate flower opening and senescence and lead
to a short vase life (Adachi et al. 1999). During
the summer season, ‘Seiun’, which is naturally a
summer-flowering cultivar, is widely produced
(Kawada, 1987). Although it is generally believed
that ‘Seiun’ is suitable for cut flowers during hot
seasons, the characteristics of cut ‘Seiun’ flowers
under high temperatures have not been investi-
gated. This study aimed (1) to confirm that senes-
cence is not accelerated at high temperatures in
‘Seiun’; (2) to investigate the effects of high tem-
peratures on carbohydrate content in these plants,
and (3) to investigate the effects of restricting
carbon sources on flower opening and senescence.
2. Materials and methods
2.1. Plant materials
Cut chrysanthemum Dendranthema ×grandiflo-
rum (Ramat.) Kitamura ‘Seiun’ plants were ob-
tained from a commercial grower in Aichi
Prefecture. The plants were cut to a length of
approximately 80 cm on 3rd August and were
immediately boxed and transported to our labora-
tory at the University of Tokyo. When they ar-
rived on 4th August, the stems were cut to a
length of 60 cm and placed in a 200-ml glass jar
containing 100 ml of deionised water. The cut
plants were divided into three groups of 25 shoots
each and placed in a growth room controlled at
20, 25, or 30°C (60 !70% RH). Another group of
plants with 20-cm stems were also placed at 25°C
(25°C/20 cm). The light intensity was constantly
adjusted to 30 !mol m − 2 s − 1 by metalhalide
lamps (Yoko lamp, Toshiba) at the top of the
shoots since changes in light conditions will affect
photosynthesis and carbohydrate analysis. The
shoots were placed 15× 20 cm apart and samples
were collected at 0, 2, 4, 8, 16 and 26 days after
the start of treatment. Five shoots were randomly
selected from each room based on a table of
random numbers.
2.2. De!elopment
To provide indices for the three-dimensional
development and wilting of the capitula, the di-
ameter (Fig. 1a) and the angle of the florets (Fig.
1b) were measured. The diameter was determined
by taking an average of the longest diameter of
the capitula and the diameter vertical to this
longest diameter (Fig. 1). The angle of the florets
was based on an average of the four angles be-
tween a line passing through the edge of one of
the four outermost ray florets and the centre
point of the involucre and a vertical line (Fig. 1).
When the angle of the florets reached 90°, the
capitula were regarded as fully opened. When the
petals started to wilt, the edges fell downwards
and the angle of the florets increased notably.
When the petals wilted after the capitula fully
opened, the angle increased to greater than 90°.
This, the angle of the florets was considered to be
one of indicators of wilting in the capitula.
The numbers of well-developed ray florets with
petals greater than 1 cm in length, less-developed
ray florets with petals less than 1 cm in length,
and tubular-shaped florets with petals never de-
veloped, were counted at 10, 18, and 27 days.
The tubular-shaped florets could contain both
real tubular florets and ray florets since this could
not be distinguished by appearance.
2.3. Weight measurements
The plants were divided into capitula, upper
leaves (about five to six leaves), middle leaves
(about four leaves), upper stems, middle stems
and lower stems parts, and the fresh weight (FW)
of each part was measured. Involucres were not
used for the measurements since they were too
small in volume and weight. The parts were then
freeze-dried, and the dry weight (DW) was
measured.
2.4. Carbohydrate analysis
The dried samples (0.1 g) were extracted in 80%
ethanol at 80°C for 2 h, dried under vacuum, and
then dissolved in water. Dichloromethane was
added to the solution in order to remove any fatty
acids. The supernatant was used for mea-
surements of the concentrations of fructose, glu-
cose, and sucrose using an F-kit (Boehringher
 M. Adachi et al. / Posthar!est Biology and Technology 20 (2000) 63–70 65

Fig. 1. Changes in diameter (a) and angle of florets (b) in capitula of cut chrysanthemum ‘Seiun’ plants placed at 20°C/60 cm stem
length (!), 25°C/60 cm ("), 30°C/60 cm (#) and 25°C/20 cm ($). The bars represent SEs, n = 5. Average of the longest diameter
(D1) and the diameter vertical to the longest one (D2) were measured. Four angles between a line passing through the edge of the
outermost floret (f1, f2, f3, or f4) and the centre point of the involucre and a vertical line (!1) were measured and averaged.

Mannheim). The residue was used for the mea-
surement of starch with the F-kit. Fructans in the
supernatant were completely hydrolysed to fruc-
tose and glucose by 0.1 M sulphuric acid for 20
min at 100°C (Kennedy and White, 1983; Trusty
and Miller, 1991), followed by neutralisation with
0.1 M sodium carbonate. Concentrations of the
hydrolysed fructose and glucose were measured
by the same enzymic method as mentioned above.
The concentration of fructans was calculated by
subtracting the concentrations of fructose and
glucose before the hydrolysis from those after
hydrolysis.
2.5. Statistical analysis
Statistical analysis was carried out between the
four different treatments (20°C/60 cm, 25°C/60
cm, 30°C/60 cm and 25°C/20 cm), and between
days, by a one-way analysis of variance (P" 0.05)
and Duncan’s multiple range test (P"0.05). A
t-test was used for comparison between two treat-
ments or between 2 days.
3. Results and discussion
3.1. Effects of high temperature on changes in the
capitula
High temperatures suppressed development but
did not accelerate senescence in the capitula. The
diameter (Fig. 1a), the angle of the florets (Fig.
1b), FW (Fig. 2a), DW (Fig. 2b) and fructose
(Fig. 2c) and glucose (Fig. 2d) concentrations
increased more slowly at 30°C/60 cm than at
20°C/60 cm and 25°C/60 cm, until the capitula
slightly wilted at 26 days. The number of well-de-
veloped florets was significantly smaller, but the
number of tubular-shaped florets was significantly
66 M. Adachi et al. / Posthar!est Biology and Technology 20 (2000) 63–70

Fig. 2. Changes in fresh weight (a), dry weight (b), fructose (c) and glucose (d) concentrations in capitula of cut chrysanthemum
‘Seiun’ plants placed at 20°C/60 cm (!), 25°C/60 cm ("), 30°C/60 cm (#) and 25°C/20 cm ($). The bars represent SEs, n =5.

larger at 30°C than at 20°C (Table 1). It can be
assumed that high temperatures would restrict
development in the petals of the ray florets, since
Fukai et al. (1997) have reported that there are
5 – 10 tubular florets that develop during corolla
formation and that this number does not change
in chrysanthemums. A carbohydrate supply is nec-
essary for expansion in cut flowers (Rogers, 1973).
The shortage of carbohydrates can possibly be
attributed to a suppression of the development of
the capitula.
High temperature accelerates respiratory activ-
ity and the consumption of water and carbohy-
drates in cut flowers (Siegelman et al., 1958;
Coorts, 1973). The slight decrease in sugars from
4 to 8 days observed only at 30°C/60 cm (Fig.
2c,d) may be due to a slight respiratory accelera-
tion at high temperatures during this period.
Mayak and Halevy (1980) have reported that the
start of the senescent phase in cut flowers is
characterised by a decrease in FW, DW and
sugars following increases that occur during the
pre-senescent stage. In ‘Shuho-no-chikara’ cut
flowers, high temperatures trigger rapid increases
and then subsequent decreases in FW, DW and
sugar content and promote rapid senescence in the
capitula (Adachi et al., 1999). High temperatures
can induce climacteric respiration in ‘Shuho-no-
chikara’ flowers. On the other hand, high temper-
atures help to maintain water content and
carbohydrates in the capitula in ‘Seiun’ flowers,
suggesting that a rapid respiratory rise and con-
sumption of carbohydrates does not occur. ‘Seiun’
flowers might show non-climacteric respiration at
both high and moderate temperatures. Wu et al.
(1989) have reported that ‘White Sim’ cut carna-
tions show a climacteric rise and a short-vase life,
while ‘Sandra’ flowers show a non-climacteric rise
and a long vase life.
Sugars are considered to accelerate water up-
take in cut flowers (Marousky, 1971; Borochov et
al., 1976). The maintenance of carbohydrates may
contribute to the maintenance of water uptake and
retard wilting in the capitula in ‘Seiun’ flowers.
 M. Adachi et al. / Posthar!est Biology and Technology 20 (2000) 63–70 67

Table 1
Effects of temperature on numbers of well-developed ray florets, less-developed ray florets, tubular florets and total florets in cut
chrysanthemum ‘Seiun’ plants at 20 and 30°C

Days after start of Treatment

treatment

20°C 30°C

10 Ray florets (well-developed) 168 # 5.4 67.3# 8.6***

Ray florets (less-developed) 36 # 13.6 152.5 # 2.5**
Tubular florets 135.3 #13.5 93.3 # 10.7*
Total 330.3 #4.2 314#7.6
18 Ray florets (well-developed) 222 # 18.8 60.3# 2.8***
Ray florets (less-developed) 85 # 17.9 156 #4.3*
Tubular florets 0#0 126.3 # 6.1
Total 307# 31.8 343 #0.6
27 Ray florets (well-developed) 157.5#40.2 94.3# 32.7a
Ray florets (less-developed) 131.5#10.5 112 #3.5a
Tubular florets 0 #0 111.5 #3.6
Total 289 #39.9 318.3 #4.4

* Significant differences between treatments at P = 0.1.

** Significant differences between treatments at P = 0.01.
*** Significant differences between treatments at P = 0.001.
a
Non significance, n.s.

3.2. Effects of short-stem treatment on changes in
the capitula
The short-stem treatment also suppressed devel-
opment in the capitula but did not accelerate
senescence. The diameter (Fig. 1a), the angle of
the florets (Fig. 1b), FW (Fig. 2a), DW (Fig. 2b)
and fructose (Fig. 2c) and glucose concentrations
(Fig. 2d) increased more slowly at 25°C/20 cm
than at 25°C/60 cm. Because the short-stem treat-
ment reduced the number of leaves and stems, the
restriction of a carbohydrate source could be
linked to the suppression in the capitula but might
not be related to the senescence. In the short-stem
treatment, fructose (Fig. 3a) and glucose (Fig. 3b)
concentrations in the upper leaves, and the con-
centrations of fructose (Fig. 4a), glucose (Fig. 4b)
and fructans (Fig. 5a) in the upper stems, notably
decreased and then increased. These results indi-
cate that sugars in the leaves and stems were
mostly serving as a carbohydrate source for the
capitula, but that this supply was insufficient for
full development to occur.
The total DW in the cut plants increased during
all treatments, suggesting that carbohydrates
could be produced during the treatments. Mor
and Halevy (1979), Emmett (1987) and Mc-
Conchie et al. (1991) suggested that photosyn-
thates were produced in leaves and supplied
flower parts in cut flowers. In the present study,
yellowing in the leaves was not observed in any
treatments. The increase in the concentrations of
sugars in the capitula, leaves and stems during the
latter stages of the short-stem treatments might
have resulted from increased supply of photo-
synthates.
3.3. Effects of high temperature on changes in the
lea!es
High temperature triggered earlier wilting in the
leaves than did lower temperatures. FW in the
leaves initially decreased, especially at 30°C/60 cm
and then remained constant until the end of the
treatment (data not shown), suggesting that
68 M. Adachi et al. / Posthar!est Biology and Technology 20 (2000) 63–70

Fig. 3. Changes in fructose (a,c), glucose (b,d) concentrations in upper (a,b) and middle (c,d) leaves of cut chrysanthemums ‘Seiun’
plants placed at 20°C/60 cm (!), 25°C/60 cm ("), 30°C/60 cm (#) and 25°C/20 cm ($). The bars represent SEs, n = 5.

high temperatures promote water loss in the
leaves during the earlier stages of treatment.
Patterns in the concentration changes for fruc-
tose (Fig. 3c), glucose (Fig. 3d) and sucrose in the
upper leaves at 30°C were different from those at
20°C/60 cm and 25°C/60 cm. Considering that
fructose and glucose concentrations were similar
in the leaves, these concentrations in the upper
leaves could be converted from those of sucrose
by invertase during the treatment. High tempera-
tures can change the levels of hydrolytic enzy-
matic activity and change the proportion of
sugars (Markus, et al., 1981; Yamane et al., 1991;
Bancal and Triboi, 1993), resulting in changes in
the carbohydrate content. Our results also suggest
that fructose and glucose could be the main
translocating sugars, since very little sucrose was
in the capitula and stems in ‘Seiun’.
High temperatures can affect photosynthesis
due to changes in the enzymic activity (Markus, et
al., 1981; Wolf, et al., 1990), and in the present
study, water loss could reduce the carbohydrate
supply of the capitula if there was photosynthesis
occurring in ‘Seiun’ leaves during the treatments.
The slower increase in sugar concentrations at
30°C/60 cm than at 20°C/60 cm and 25°C/60 cm
in the middle leaves (Fig. 3b,d) may have been
due to suppression of photosynthetic activity by
high temperatures.
3.4. Effects of high temperature on changes in the
stems
The sum of the concentrations of fructose (Fig.
4a,c), glucose (Fig. 4b,d) and fructans (Fig. 5)
increased for all treatments; they did, however,
increase more slowly at 30°C/60 cm than at 20°C/
60 cm and 25°C/60 cm. The increase in carbohy-
drate concentrations is likely to be due to supply
from the leaves if there was no photosynthetic
production in the stems. The concentrations of
fructans in the upper (Fig. 5a), middle (Fig. 5b)
and lower (Fig. 5c) stems increased at 20°C/60 cm
and 25°C/60 cm, suggesting that the stems were
being used for the storage of sugars.
 M. Adachi et al. / Posthar!est Biology and Technology 20 (2000) 63–70 69

Fig. 4. Changes in fructose (a,c) and glucose concentrations (b,d) in upper (a,b) and middle (c,d) stems of cut chrysanthemum ‘Seiun’
plants placed at 20°C/60 cm (!), 25°C/60 cm ("), 30°C/60 cm (#) and 25°C/20 cm ($). The bars represent SEs, n= 5.

High temperatures appeared to reduce the sup-
ply from leaves or accelerate the consumption of
carbohydrates as respiratory substances in the
stems as well as the capitula. In addition, high
temperatures appeared to accelerate the hydroly-
sis of fructans and increase fructose (Fig. 4a,c)
and glucose (Fig. 4b,d) concentrations in the
stems. In ‘Shuho-no-chikara’ cut flowers, sugars
in the stems notably decrease under these condi-
tions, particularly, in the middle stems, suggesting
that these could be a source for the capitula
(Adachi et al., 1999). Middle stems might also be
a source for the capitula at moderate tempera-
tures in ‘Seiun’ cut flowers.
In summary, high temperatures and short-stem
treatments suppress development but do not ac-
celerate senescence in the capitula. A carbohy-
drate supply might be necessary for the
development of the capitula. High temperatures
and the short-stem treatment could not rapidly
reduce the water and carbohydrate content in the
capitula. In addition, respiration and the con-
sumption of carbohydrates do not appear to
change significantly during the treatments, result-
ing in the maintenance of the carbohydrate and
the retarding of senescence. Because cut chrysan-
themums are sold before the capitula are fully
opened, high temperatures after harvest may re-
sult in poor development of the capitula, and
subsequent loss of quality.
Acknowledgements
The authors wish to thank Dr Mikihiro Hara at
the Floricultural Institute, Aichi Agricultural Re-
search Institution for generously providing cut
chrysanthemum plants for this study, and Dr
Anthony D. Kendle in the Department of Horti-
culture, University of Reading, UK, for his care-
ful reading of the manuscript.
70 M. Adachi et al. / Posthar!est Biology and Technology 20 (2000) 63–70
Fig. 5. Changes in concentration of fructans in upper (a),
middle (b) and lower (c) stems of cut chrysanthemum ‘Seiun’
plants placed at 20°C/60 cm (!), 25°C/60 cm ("), 30°C/60 cm
(#) and 25°C/20 cm ($). The bars represent SEs, n = 5.
References
Adachi, M., Kawabata, S., Sakiyama, R., 1999. Changes in
carbohydrate content in cut chrysanthemum (Den-
dranthema×grandiflorum (Ramat.) Kitamura ‘Shuho-no-
chikara’) plants during flower opening and senescence at
different temperatures. J. Jpn. Soc. Hort. Sci. 68, 228–235.
Bancal, P., Triboi, E., 1993. Temperature effect on fructan
oligomer contents and fructan-related enzyme activities in
stems of wheat (Triticum aesti!um L.) during grain filling.
New Phytol. 67, 517 –531.
Borochov, A., Tirosh, T., Halevy, A.H., 1976. Abscisic acid
content of senescing petals on cut rose flowers as affected
by sucrose and water stress. Plant Physiol. 58, 175–178.
Cockshull, K.E., Kofranek, A.M., 1994. High night tempera-
tures delay flowers, produce abnormal flowers and retard
stem growth of cut flower chrysanthemum. Sci. Hort. 56,
217 – 234.
Coorts, G., 1973. Internal metabolic changes in cut flowers.
HortScience 8, 195 – 198.
Emmett, M.R. 1987. Improving chrysanthemum vase life.
ADAS Horticulture Services, London, p. 23.
Fukai, S., Zhang, W., Uehara, H., Goi, M., 1997. Morpholog-
ical changes in shoot apex during floral initiation and
development in chrysanthemum (Dendrathema grandiflorum
(Ramt.) Kitam.). Tech. Bull. Fac. Agr. Kagawa Univ. 49,
171 – 178.
Funakoshi, K., 1984. Studies on the influence of growing and
environmental conditions on the morphological characteris-
tics and vase life of cut flowers in Chrysanthemum mori-
folium Ram. Special report 15th Shizuoka Agricultural
Institution.
Kawada, J., 1987. The phasic development of chrysanthemum
as a basis for the regulation of vegetative growth and
flowering in Japan. Acta Hort. 197, 115 – 123.
Kennedy, J.F., White, C.A., 1983. Bioactive carbohydrates in
chemistry, biochemistry and biology. Halsted Press,
Chuchester, New York.
Markus, V., Lurie, S., Bravdo, B., Stevens, M.A., Rudich, J.,
1981. High temperature effect on RuBP carboxylase and
carbonic anhydrase activity in two tomato cultivars. Phys-
iol. Plant. 53, 407 – 412.
Marousky, F.J., 1971. Inhibition of vascular blockage and
increased moisture retention in cut roses induced by pH,
8-HQC and sucrose. J. Am. Soc. Hort. Sci. 96, 38 – 41.
Mayak, S. Halevy, A.H., 1980. Flower senescence. In: Thi-
mann, K.W. (Ed.), Senescence in Petals, CRC Press, Boca
Raton, Florida, pp.131 – 156.
McConchie, R., Lang, N.S., Gross, K.C., 1991. Carbohydrate
depletion and leaf blacking in Protea neriifolia. J. Am. Soc.
Hort. Sci. 116, 1019 – 1024.
Mor, Y., Halevy, A.H., 1979. Translocation of 14C-assimilates
in roses. II. The effect of the shoot and the location of the
source leaf. Physiol. Plant. 45, 177 – 182.
Rogers, M.N., 1973. An historical and critical review of
postharvest physiology research on cut flowers. HortScience
8, 189 – 194.
Siegelman, H., Chow, C.T., Biale, J.B., 1958. Respiration of
developing rose petals. Plant Physiol. 33, 403 – 409.
Trusty, S.E., Miller, W.B., 1991. Postproduction carbohydrate
levels in pot chrysanthemums. J. Am. Soc. Hort. Sci. 116,
1013 – 1018.
Whealy, C.A., Nell, T.A., Barrett, J.E., 1987. High temperature
effects on growth and floral development of chrysanthe-
mum. J. Am. Soc. Hort. Sci. 112, 464 – 468.
Wolf, S., Olesinski, A.A., Rudich, J., Marani, A., 1990. Effect
of high temperature on photosynthesis in potatoes. Ann.
Bot. 65, 179 – 185.
Wu, M.J., van Doorn, W., Mayak, S., Reid, M.S., 1989.
Senescence of ‘Sandra’ carnation. Acta Hort. 261, 221 – 225.
Yamane, K., Kawabata, S., Sakiyama, R., 1991. Change in
water relations, carbohydrate contents, and acid invertase
activity associated with perianth elongation during anthesis
of cut gladiolus flowers. J. Jpn. Soc. Hort. Sci. 60, 421 – 428.