Journal of Horticultural Science

ISSN: 0022-1589 (Print) (Online) Journal homepage: http://www.tandfonline.com/loi/thsb19

Effects of Ethylene on Carnation Flowers

(Dianthus Caryophyllus) Cut at Different Stages of
Development

P. Camprubi & R. Nichols

To cite this article: P. Camprubi & R. Nichols (1978) Effects of Ethylene on Carnation Flowers
(Dianthus Caryophyllus) Cut at Different Stages of Development, Journal of Horticultural Science,
53:1, 17-22, DOI: 10.1080/00221589.1978.11514788

To link to this article: https://doi.org/10.1080/00221589.1978.11514788

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Journal of Horticultural Science (1978) 53 (17-22)

Effects of ethylene on carnation flowers

(Dianthus caryophyllus) cut at different stages of development
By P. CAMPRUBI* and R. NICHOLS
Glasshouse Crops Research Institute, Littlehampton, W. Sussex BN 16 3PU, UK

SUMMARY
Carnation flowers cut at various stages of development became increasingly sensitive to
ethylene as they aged. Ethylene (e.g. 1 vpm for 24 h) caused irreversible wilting of open
flowers but had little or no effect on the youngest flowers (buds), provided they were
opened in a sucrose-germicide solution. The tolerance of flowers to exogenous ethylene or
propylene was inversely related to their capacity to produce endogenous ethylene. Ten vpm
ethylene for 24 h induced elongation and often an increase in the fresh weight of the petals
of the youngest flowers and, after eight days, there was enlargement of the gynoecium: in
contrast, older buds failed to open and mature flowers wilted. The implications of thM; work
are discussed in relation to the marketing of carnation flowers.

ALTHOUGH the sensitivity of carnation flowers to morphology of carnation flowers cut at different
ethylene has long been known (Crocker and stages of development.
Knight, 1908), recent studies have shown that the
maturity of the flower, temperature, and partial MATERIALS AND METHODS

pressures of 0 2 and C0 2 influence the response of Flowering shoots of D. caryophyllus cv White

the flower to ethylene (Smith and Parker, 1966;
Nichols, 1968; Uota, 1969; Maxie, Farnham,
Mitchell, Sommer, Parsons, Snyder and Rae,
1973; Dilley and Carpenter, 1975).
Barden and Hanan ( 1972) showed that im-
mature flowers (buds) were relatively tolerant of
low doses of ethylene, particularly at low
temperatures, and this could be an additional
reason tor the practice of picking and trans-
porting the flowers in bud, in which there is
increasing interest.
There is evidence that endogenous ethylene is
involved in the natural senescence of the mature
carnation flower (Nichols and Ho, 1975; Mayak
and Dilley, 1976), but the different responses of
buds and mature flowers to exogenous ethylene
(Barden and Hanan, 1972) suggest that the fac-
tors leading to their senescence may differ. The
object of this work was to compare the effects of
ethylene on some aspects of the physiology and
*Present address: C.R.I.D.A. 04, I.N.I.A., Cabrils
(Barcelona). Spain.
Sim were cut at Stages 3, 4 and 6 (Fig. I) and the
stems trimmed to 30 em. Each cut flower was
weighed; those weighing more or less than the
mean ± 1.5 x the standard deviation of the pop-
ulation mean weight were discarded. Not less
than five flowers were used in each treatment and
experiments were repeated a number of times;
pooled data are presented where practicable. All
observations were made at l8·3°C and 60% r.h.
Water uptake was measured gravimetrically
(Nichols and Ho, 1975).
Gas analyses and treatments
Ethylene and propylene were measured in a gas
chromatograph fitted with a Poropak S {Waters
Associates, Ltd) analytical column and flame-
ionization detector. Oxygen and C0 2 were
analysed in a chromatograph fitted with a
molecular sieve (5A, BDH Ltd) analytical
column and a thermistor detector assembly.
Flowers with the cut ends of their stems in
water were treated with olefins inside closed 2(}-I
jars into which the estimated volume of ethylene
or propylene was injected through a septum.
18 Effects of ethylene on carnation flowers cut at different stages
FIG. I
Stages 3, 4 and 6 of flower development.
After 1 h and just before opening the jars, 1-ml
samples of gas were withdrawn for analyses of
the gas concentrations. Beakers of NaOH (20%)
were included in the jars to absorb respiratory
C0 2 • After the olefin treatment the flowers were
transferred to water or a solution of 8% sucrose
containing a germicide, either dichlorophen (25
mg 1-1) or 8-hydroxyquinoline citrate (HQC, 200
mg I-1), to promote flower development; without
this treatment carnation flower buds may fail to
open regardless of the nature of the gas
treatment. To measure the effects of olefin
treatment on endogenous ethylene production,
single flowers were put in respiration jars fitted
with sampling ports (Nichols, 1966). Fresh and
dry weights were obtained by removing and
blotting the tissue, weighing it in tared containers
and then reweighing after oven-drying at 80°C for
24h.
RESULTS
In preliminary experiments we confirmed -that
flowers treated at Stage 6 with 1 vpm ethylene for
24 h wilted irreversibly during or shortly after the
flowers were removed from the gas treatment.
This agrees broadly with the reports of the sen-
sitivity of mature flowers to ethylene (Barden and
Hanan, 1972; Maxie et a/., 1973; Nichols and
Ho, 1975).
To verify that 0 2 was not depleted to levels
that would cause anaerobiosis during gas
treatments, 0 2 concentrations inside the jars were
measured at the end of the 24-h treatment on four
occasions. The mean 0 2 levels in jars containing
Stage 3 flowers and 0 or 10 vpm ethylene were
15·7% (range 15·2 to 16·3%) and 15·1%
(14·4-16-2%) respectively. Most of the respired
C0 2 was absorbed by the NaOH, and the con-
centrations, sampled at the tops of the jars, varied
between 0·08 and 0·18% depending on the
numbers of flowers put in the jars.
Tables lA and IB show the survival of Stage 3,
4 and 6 flowers after treatment with one of a
range of ethylene concentrations; comparisons
between these tables are not strictly valid since
the experiments were not conducted at the same
time and there appears to be a seasonal or pre-
harvest environment effect (q.v.) on sensitivity.
It is evident that bud-cut flowers tolerate doses
of ethylene (0·5-1·0 vpm for 24 h) which are
toxic to Stage 6 flowers. Clearly there was an
increasing sensitivity to ethylene as the buds
developed - Stage 4 buds, unlike those at Stage 3,
did not open in concentrations of ethylene as high
as 10 vpm.
During these experiments we observed that
petals of the 10 vpm ethylene-treated Stage 3
flowers enlarged slightly, more than the untreated
controls. Since this was unexpected, because
ethylene generally inhibits cell elongation, possi-
ble causes were investigated.
 P. CAMPRUBI and R. NICHOLS 19

Effect of ethylene treatment on petals and water
uptake of Stage 3 and 4 flowers
Ethylene promoted water absorption and, in
Stage 3 flowers, an enlargement of the petals
(Table IIA and liB). The Stage 4 flowers failed to
open in I 0 vpm ethylene, and so these data have
been omitted from Table liB. In these ex-
periments absorbed water was retained by stem,
leaves and corolla since the whole flowering stem
was treated; there would have been little trans-
piration in the closed jars of the gas treatment
system. However, it was not possible to confirm
that the increased petal length (Table liB) was ac-
companied by increases in their fresh weight, if all
the experimental data were combined as in Table
liB. In the first six of eight experiments, con-
ducted in spring and early summer, ethylene
promoted a small but significant increase in fresh
weight (P=0·05), whereas in the last two, in mid-
summer, the petal weights just failed to differ
significantly between the treated and untreated
flowers.
A possible criticism is that in Stage 3 flowers
the petals are enclosed to a large extent inside the
calyx whereas they are exposed in Stage 6
flowers. However, since the Stage 3 flowers show
an elongation response it is evident that the
ethylene penetrates the tightly furled corolla.

TABLE lA
Effects of ethylene (24 h treatment) on longevity (days) of Stage 3 and 4 flowers. After the gas treatment the flowers were put
in sucrose-dichlorophen until open (Stage 6) and then transferred to water.
Time to open to Ethylene concentration (vpm)
Flower stage at
Stage 6
gas treatment
(days) 0 0·5 1·0 10·0
3 15 5·0(0·4) 5·1(0·6) 4·9(0·4)
4 7 6·0(0·5) 5·8(1·0) 4·8(1·0) 0
SE of mean in parentheses; n=not less than 15 per treatment.

TABLE 18
Effect of ethylene on longevity (days) of Stage 4 and 6flowers; details as in Table !A
Ethylene concentration (vpm)
Flower stage at
gas treatment
0 0·2 0·5

4 7·2(0·4) 7· 7(0·2) 7·0(0·4)

6 7·5(0·2) 6·9(0·1) 3·0(0)

TABLEIIA
Mean water uptake (gflower-• d- 1) of Stage 3 and 4 flowers after ethylene treatment for 24 h.
Ethylene concentration (vpm)
Stage
0 10
3 1·26"(0·03) 1·40tx0·05) 1·53b(0·06)
4 ]. 74C(0·06) 2·32.(0·10) 2 ·13.(0·08)

The means followed by the same letter are not significantly different (P=0·05); SE of the mean in parentheses;
n=25 per treatment.

TABLE liB
Mean fresh weight and length of petals from Stage 3 flowers after ethylene treatment for 24 h.
Fresh weight Length
Ethylene concentration (vpm)
(g) (em)

0 3·16 4-01
10 3·31 4·37
LSD (P=0·05) 0·16 0·11

n for fresh weight=30 flowers; n for length=15 flowers, 3 petals per flower.
20 Effects of ethylene on carnation flowers cut at different stages

Comparison of endogenous ethylene production
in Stage 3 and 4 flowers
We were unable to detect stimulation of en-
dogenous ethylene in Stage 3 flowers, after expos-
ing them to I or I 0 vpm ethylene for 24 h. The
smaller dose caused only a small amount to be
produced from Stage 4 flowers, but it was suf-
ficient to cause petal wilting in Stage 6 flowers
(Table IB) and enhanced ethylene production.
Larger doses of ethylene (I 0 vpm), or roughly
analogous amounts of propylene ( I500 vpm,
Burg and Burg, I965) induced endogenous
ethylene in Stage 4 flowers. A typical time-course
of ethylene evolution from a gas-treated Stage 4
flower is shown in Table III. It seems that one
difference between the flower stages with regard
to their sensitivity to ethylene is their capacity to
produce endogenous ethylene.
Effect of the pre- and post-harvest environment on
the ethylene response
Examination of the data from a number of ex-
periments suggested that there was an effect of
the pre-harvest environment on the sensitivity of
the buds to ethylene; this is indicated by the data
and sucrose treatments. The main effect (Table
V) was the expected increase in dry weights of
petals and gynoecium caused by the sucrose
treatment. However, there appeared to be a small
but significant increase in the gynoecium caused
by the ethylene treatment above that caused by
the sucrose alone. The promotion of ovary
growth by ethylene has been described for Stage
6 flowers (Nichols and Ho, I975) and was
usually, but not exclusively (Nichols, I97I),
associated with doses of ethylene which caused
petal wilting; in these current experiments with
Stage 3 flowers the promotion of ovary growth
occurred without petal wilting.
DISCUSSION
In this paper we have shown that there is an
increasing sensitivity to exogenous ethylene as
carnation flower buds age. Doses of ethylene
which cause irreversible wilting of mature (Stage
6) flowers may have no effect on the survival and
display life of buds subsequently opened in a
sugar/germicide solution. Broadly, these obser-
vations confirm those of Barden and Hanan
(I 972); although there are evident differences in

TABLE III
Ethylene production (nlf/ower-• h-1) by Stage 4 flowers treated with ethylene or propylene
Gas treatment

Concentration Time
(vpm) (h) Time after gas treatment (h)
propylene ethylene 31 48 72 98 168

0 24 0 0 0 0 0
I 24 0 20 24 23 26
10 24 113 461 612 (w)
I 48 132 471 808 (w)
1500 24 0 45 100 109 250 (w)

w=flowers visibly wilting

in Table IV for Stage 4 buds treated with I·O vpm TABLE IV

for 24 h. We did not find the same variability in Effect ofpre-harvest environment and season on
longevity (days) of Stage 4 buds in response to ethylene
response with Stage 3 buds which were tolerant (14 h); detail as Table /A.
of this dose of ethylene.
In five separate experiments the residual effect Ethylene concentration (vpm)
Samphng date
of ethylene (I 0 vpm for 24 h) on the fresh and dry 0 1.0
weights of petals, stems, leaves and gynoecium of 15.iii.76 7 6
5.iv.76 8 8
Stage 3 flowers was measured on day 9. The 17.v.76 7 0
stems were placed in either sucrose (8%) contain- 29.v.76 7 0
8.xi.76 7 6
ing HQC (200 mg 1- 1) or distilled water to find
out if there are interactions between the ethylene n=5 flowers per sampling date
 P. CAMPRUBI and R. NICHOLS 2I
TABLEV
Mean fresh and dry weights of flowers treated with ethylene as Stage 3 buds on day 9. After the gas treatment (24 h) the
flowers were transferred to water or sucrose-HQC.
Petals Stem Leavest Gynoecium
Ethylene
Solution treatment FW FW DW FW BW FW DW
(vpm) DW
(g) (g) (g) (g) (g) (g) (g) (g)

Water 0 3·84 0·69 4·30 0·73 0·256 0·048 0·2252 0-0352

10 3-88 0·60 4·35 0·73 0·243 0·046 0·2650 0·0393
Sucrose·HQC 0 4-80 0·94 4-35 0·98 0·271 0·055 0-3157 0-0596
10 5·51 0·93 4·51 1·02 0·255 0·054 0·3699 0-0683
LSD (P=0·05) 0-30 0·05 0·31 0·09 0·031 0-006 0-0264 0-0063

t for mean wt per leaf, n=25.

detail, they are not surprising in view of the
differences in techniques, plant material and the
pre- and post-harvest environments. Since we
used two bud stages we were able to show that
there is no sharp change in the morphological
stage at which the flowers become more sensitive
to ·ethylene. The visible response to I vpm for 24 h
ethylene with regard to petal wilting was in the
sequence:
Stage 3 < Stage 4 < Stage 6
This dose of ethylene was chosen for its clear
effect on the Stage 6 flowers; it is also a realistic
level in commercial terms. At higher doses the
responses of Stage 4 and 6 flowers were in-
distinguishable- they both wilted irreversibly, the
former as buds without opening, the latter as
mature flowers. From these observations we con-
clude that there is a progressive change in the
physiology of the ethylene response as the flower
ages. This is recognized in fruit tissues which re-
quire smaller doses of ethylene to enhance ripen-
ing as the fruit ages and approaches ripening
(Abeles, I973).
A possible explanation for the change in sen-
sitivity of buds to ethylene is that their capacity to
produce endogenous ethylene increases as they
age. Levels of ethylene which induce rapid wilting
of Stage 6 flowers promote high levels of en-
dogenous ethylene (Nichols, 1968). In this work a
dose of ethylene (I vpm for 24 h), which caused
irreversible wilting of Stage 6 flowers, induced
only small amounts of endogenous ethylene from
Stage 4 buds (Table III), and although this dose
was not sufficient to cause immediate wilting, it
generally shortened the survival (Table lA) of
flowers opened from this stage. High doses of
ethylene or propylene induced over 100 nl h- 1 of
endogenous ethylene, which increased as the
flowers aged (Table III), and this was accom-
panied by failure of the buds to open. Thus it
seems that variation in the survival of the
ethylene-treated buds, within and between stages,
may depend on their capacity to produce en-
dogenous ethylene. For this reason it is difficult to
describe threshold doses of ethylene except in the
broadest terms. The concept of a threshold dose
includes a number of largely subjective elements
based on the recognition of flower survival and a
choice of dosage levels.
The observation that petals of Stage 3 buds
elongated in response to I 0 vpm ethylene was un-
expected (Table liB), since the same dose was
toxic to Stage 4 buds. It suggests that promotion
of tissue elongation by ethylene which has been
reported to occur, for example, in rice coleoptiles
(Ku, Suge, Rappaport and Pratt, 1970), stems of
Ca/litriche p/atycarpa (Musgrave, Jackson and
Ling, I972) and petioles of Ranunculus
sce/eratus (Musgrave and Walters, I973), also
occurs in the immature petals of carnation. The
petal elongation is likely to be a complex
metabolic interaction with endogenous auxins
and gibberellins which have been reported to oc-
cur in carnation flowers (Jeffcoat and Harris,
1972). The implication is that ethylene enters into
the metabolism of the flower bud, and there is
evidence for this in other plant tissue (Beyer,
1975). Ethylene increases respiration of mature
carnation flowers (Nichols, 1968; Maxie et ,,a/.,
1973). In these, the ethylene-enhanced respiration
accompanies or precedes death of the petals, but
little is known of the physiological effects of "sub-
lethal" doses of ethylene on mature carnation
flowers, which can show partial loss of turgor and
then recovery, following low doses of ethylene
(Nichols, 1968; Mayak and Dilley, 1976).
If we consider the horticultural implications of
this work we find that Stage 3 buds take too long
to open (Table lA) although these were the most
22 Effects of ethylene on carnation flowers cut at different stages

ethylene-tolerant of the buds that we worked
with; Stage 4 buds are the youngest that could be
opened in a commercially acceptable time. The
sucrose-germicide solution may not have been
optimal for opening these buds but it was
appropriate for the needs of this work.
Nonetheless, Stage 4 buds will tolerate doses of
ethylene in excess of those likely to be met in nor-
mal commercial UK marketing, particularly if the
interaction with temperature is considered.
This work was done at the Glasshouse Crops
Research Institute whilst P.C. was the recipient of
a World Bank - INIA Fellowship.

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