Postharvest Biology and Technology 21 (2001) 303– 310

www.elsevier.com/locate/postharvbio

Benzyladenine and the vase life of tropical ornamentals

Robert E. Paull a,*, Theeranuch Chantrachit b
a
Department of Tropical Plant and Soil Sciences, College of Tropical Agriculture and Human Resources,
Uni6ersity of Hawaii at Manoa, 3190 Maile Way, Honolulu, HI 96822, USA
b
Faculty of Agricultural Production, Maejo Uni6ersity, Sansai, Chiang Mai 50290, Thailand

Received 8 July 1999; accepted 29 July 2000

Abstract

The vase life of anthurium (Anthurium andraeanum), Heliconia (Heliconia psittacorum cv. ‘Andromeda’, H.
chartacea cv. ‘Sexy Pink’), red and pink ginger inflorescence (Alpinia purpurata) was increased by benzyladenine (BA,
100 mg L − 1), applied as a dip or as a spray. There was no effect of BA on the vase life of Bird of Paradise (Strelitzia
reginae), Beehive ginger (Zingiber spectabilis), and Uluhe fern curls (Dicranopteris linearis). Different anthurium
cultivars showed differences vase life response to BA treatment from a 20% reduction to a 2.5 fold increase in vase
life. Anthurium cultivars that responded positively to BA and were packed for 8 days had 20 days longer vase life
than non-BA treated flowers. The effects of BA on topical foliage varied, while the vase life of anthurium leaves was
extended by BA and that of the flowerless Bamboo orchid (Arundina bambusifolia) and the fern Lycopodium
(Lycopodium cernuum) leaves were shortened. The data suggested that the effect of BA on the extension of vase life
depended upon flower or inflorescence type, season of harvest and cultivar. © 2001 Elsevier Science B.V. All rights
reserved.

Keywords: Cut flowers; Foliage; Packing; Storage temperature; Application time

1. Introduction Ginger (Alpinia purpurata) and Heliconia (Helico-

Tropical flowers frequently have insignificant
inflorescences and flowers, and the bracts are
colorful and give the ‘flower’ its appeal. The
anthurium (Anthurium andraeanum) flower con-
sists of a modified leaf; the spathe, and a flower
spadix with more than 300 spirally attached
minute flowers (Watson and Shirakawa, 1967).
* Corresponding author. Tel.: +1-808-9567369; fax: +1-
808-9563894.
E-mail address: paull@hawaii.edu (R.E. Paull).
nia psittacorum) are similar having bracts
(modified leaves) and a few conspicuous flowers.
The vase life of these flowers can vary greatly, for
example, anthurium vase life seasonally can vary
from 8 to 69 days (Paull et al., 1992) and different
Heliconia varieties from 7 to 21 days (Powell,
1989).
The cytokinin, benzyladenine (BA), increases
the vase life of anthurium, Heliconia and red
ginger from 1.2 to 1.7-fold, while Bird of Paradise
shows no response to BA (Whittaker, 1993).
Other flowers, such as carnations (MacClean and

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PII: S 0 9 2 5 - 5 2 1 4 ( 0 0 ) 0 0 1 5 3 - 8
304 R.E. Paull, T. Chantrachit / Posthar6est Biology and Technology 21 (2001) 303–310
Dedolph, 1962), iris, rose, tulips and daffodils,
show a slight positive effect or no increase in vase
life in response to BA application (cf. Halevy and
Mayak, 1979). Whittaker (1993) dipped an-
thurium, Heliconia and red ginger in 200 mg L − 1
BA dips after the flowers were shipped from
Jamaica to Florida. In earlier work (Shirakawa et
al., 1964), momentarily dipped anthuriums stems
into a solution containing 10 mg L − 1 BA plus
0.1% Tween-20, after 30 h shipping from Hawaii
to Michigan, and the flowers showed only a mar-
ginal increase in vase life. The addition of benzyl-
riboside to the vase solution increases the vase life
of ‘Ozaki’ anthurium, while benzyladenine in the
vase solution did not increase the vase life in
preliminary experiments (Paull, 1984, Unpub-
lished). The differences in anthurium results could
be due to cultivar, the delay between harvest,
shipment and treatment, and, the concentration of
BA used. Whittaker (1993) used the anthurium
cultivars, ‘Red Obake’, ‘Bettina’ and ‘Nitta’, and
found cultivar differences in the degree of re-
sponse to BA. The anthurium ‘Ozaki’ was used by
Paull (1984, Unpublished), while Shirakawa et al.,
(1964) did not indicate the cultivars used. Another
possible explanation for the difference in results is
that both Whittaker (1993) and Shirakawa et al.,
(1964) treated their flowers after a period of pack-
ing and shipping of up to 30 h. Their postharvest
handling sequence was to harvest, pack, ship,
unpack then a BA dip, while Paull (1984, Unpub-
lished) applied the BA in the vase solution. The
objective of the reported series of experiments was
to determine the response of a range of tropical
flowers and foliage to BA application applied
prior to and after packing and simulated shipping.
2. Materials and methods
2.1. Flower supply
Flowers and foliage were obtained on the day
of harvest from commercial sources in Hilo,
Hawaii. After BA treatment and packing in Hilo,
the cartons were shipped to Honolulu that after-
noon. Red ginger flowers were grown at Poamoho
Experiment Station on Oahu, harvested, held in
water and transported to the laboratory within 2
h.
Anthurium flowers had at least three-quarters
open flowers on the spadix, unless indicated oth-
erwise. Anthuriums were discarded if they showed
any spathe or spadix damage.‘Andromeda’ and
‘Sexy Pink’ Heliconia were harvested when the
first bract had opened. Red and pink ginger infl-
orescences were two-thirds open when harvested.
Bird of Paradise inflorescences were harvested
when one floret had emerged, and beehive gingers
when the bracts had fully opened. The foliage
material used was the fern Lycopodium, the flow-
erless Bamboo Orchid, and anthurium leaf.
2.2. Application method and handling
The flowers and foliage were dipped by sub-
merging for 10 s to 1 min, depending on the
experiment, or, alternatively, sprayed with a fine
mist to cover all surfaces of the flower and foliage.
Routinely, 200 mg L − 1 BA was routinely used,
following the results of Whittaker (1993). In two
anthurium experiments, BA was sprayed onto the
flowers either before or after packing. After BA
treatment, flowers and foliage were allowed to
dry, then packed in moistened, shredded newspa-
per in standard flower boxes with a plastic liner.
The cartons were normally held for 2 days at
22°C, and in additional series of experiments up
to 12 days at 8°C or 22°C. Upon unpacking, 4–5
cm of stem was removed and flowers were placed
in 1 L flasks of deionized water. Vase life was
evaluated at 22°C, 70–80% RH, four air changes
per hour in a room receiving fluorescent light (10
W m − 2) for 12 h per day. Vase deionized water
was replenished every week.
2.3. Flower and foliage e6aluation
Flowers were evaluated individually and vase
life was measured as the time till the flowers and
foliage were discarded. Anthurium spadix senes-
cence (necrosis) was ranked on a scale: 1- (good,
no senescence spadix) to 5- (unacceptable, 50% or
more of spadix showing senescence); the spathe
blueing scale: 1- none to 4- 100% of the spathe
area showing blueing, and spathe gloss scale: 1-
 R.E. Paull, T. Chantrachit / Posthar6est Biology and Technology 21 (2001) 303–310 305

no gloss loss to 4- full gloss loss and wilting.
Flowers were discarded if they were rated 4 for
spadix senescence, 3 for spathe color, or 4 for
spathe gloss (Paull and Goo, 1982). Bird of Par-
adise was discarded when 50% of opened-florets
had wilted. ‘Sexy Pink’ and ‘Andromeda’ Helico-
nia were discarded when all florets in the opened
bracts had dropped, or the tip of flower bract had
turned black on 50% of the total bract area. Pink
and red gingers inflorescences were evaluated on a
senescence scale: 0 – no senescence symptoms, to,
4 – more than 50% of total bract area showed
browning, and inflorescence wilted scale: 0 – no
wilting symptoms, to, 4 – the inflorescence had
lost turgor and drooped from the vertical axis by
more than 90°. Beehive ginger was discarded
when all flowers were opened and when more
than 50% of the total bract area had turned
brown. The foliage materials, anthurium leaf, Ly-
copodium and Bamboo Orchid were discarded
when more than 50% of the total area showed
yellowing or was dried out. Uluhe fern curls were
discarded when the stem had lost turgor and more
than 50% of the stems showed drooping. All
inflorescence and foliage were evaluated every
other day after unpacking.
2.4. Data analysis
Data was subjected to analysis of variance or
regression using the GLM procedure (SAS Insti-
tute, Cary, NC). Treatment means were compared
by the Duncan –Waller multiple range tests or
LSD test at the 5% probability level.
3. Results and discussion
Different anthurium cultivars, dipped into 200
mg L − 1 BA, showed a variable effects on vase life
from a 20% reduction to a 2.5 fold increase (Table
1). Paull (1984, unpublished) used ‘Ozaki’ in the
earlier tests and the lack of response of this
cultivar to BA was confirmed in these tests, where
‘Ozaki’ consistently showed a reduction or only a
slight increase in vase life following BA treatment.
The anthurium cultivar ‘Leilani’ and ‘Marian See-
furth’ consistently showed the greatest response to
BA of more than a 2.5 fold increase in vase life.
The reason for the wide variation in cultivar
responsiveness to BA was unclear as it does not
apparently relate to the parent lines used to de-
velop these cultivars (Kamemoto, 1997, personal
communication).

Table 1
Effect of dipping different Anthurium cultivars in BA (200 mg L−1) on flower vase life. Flowers were packed in a carton for 2 days
and held at 22°C before unpacking, 4–5 cm cut from stem and placed in deionized water for vase life evaluation

Cultivars Vase Life (days from harvest)a

No BA treatment Treated BA Increase vase life times (×)

Blushing Bride 31 b 44 a 1.5

Blush Oishi 26 b 40 a 1.5
Kalapana 14 b 32 a 2.3
Kozohara 29 b 42 a 1.4
Lavender Lady 15 b 30 a 2.0
Marian Seefurth 19 b 46 a 2.5
Mickey Mouse 20 b 40 a 2.0
New Pahoa Red 16 b 51 a 3.2
Nitta 34 b 51 a 1.6
Oshiro Red 18 b 40 a 2.2
Oshiro White 19 b 37 a 2.0
Ozaki 24 a 19 b 0.8
Rainbow 27 b 43 a 1.6
Tatsata Pink 17 b 41 a 2.4
Tropic Mist 27 b 43 a 1.6

a
Means with same letter in a row were not significantly different by Duncan–Waller multiple range test at P50.05, n = 12.
306 R.E. Paull, T. Chantrachit / Posthar6est Biology and Technology 21 (2001) 303–310
Fig. 1. Effect of different BA concentrations on the vase life of
five anthurium cultivars: Ozaki, Purple Arc, Blush Oishi,
Marian Seefurth and Leilani, packed for 2 days unpacked,
4–5 cm cut from the stem, and placed in deionized water for
vase life evaluation. Analysis of variance: Cultivar, P= 0.0001;
BA concentration, P=0.0001; Cultivar × concentration, P=
0.0001.
There was significant variation in the pattern of
response of different cultivars to BA concentra-
tion (Fig. 1). The maximum vase life increase was
obtained at about 100 mg L − 1 BA for ‘Leilani’,
‘Blush Oishi’, and ‘Purple Arcs’. ‘Marian See-
furth’ reached the maximum vase life at 150 mg
L − 1 and gave only a slight further increase in
vase life at 200 mg L − 1. ‘Ozaki’ showed a slight
increase in vase life at 100 mg L − 1 (Fig. 1), while
10 mg L − 1 used by Shirakawa et al., (1964) gave
little or no increase in vase life. There was no
reduction in vase life at higher BA concentrations.
The significant correlation suggested that even a
slight increase in available cytokinin gave an in-
crease in vase life.
Those cultivars that responded most to BA had
either a greater capacity to absorb the applied BA
or had low tissue levels of cytokinins. The lack of
response shown by ‘Ozaki’ was not apparently
related to the absorption of BA, as inclusion of
detergent in the BA dip did not increase vase life
(data not shown). ‘Ozaki’ vase life, in this study,
was between 18 and 25 days and has been re-
ported to be up to 69 days (Paull et al., 1992) and
it lacks of response to BA may be due to high
natural levels of cytokinins. Preliminary results on
‘Leilani’ showed that BA maintained water up-
take at ca. 1.5 mL h − 1 g − 1 FW, while the un-
treated flower’s water uptake was ca. 0.5 mL h − 1
g − 1 FW. ‘Ozaki’ flowers had a higher uptake rate
(2.5 mL h − 1 g − 1 FW) but there was little differ-
ence between BA treated and untreated flowers.
This difference in water uptake may be related to
the maintenance of the spadix whose condition is
a major cause of loss of appearance (Paull and
Goo, 1985).
In initial tests, the whole anthurium flower was
dipped in BA. This treatment was then compared
to spraying just the top or spraying both the
adaxial and abaxial spathe surfaces (full spray). A
similar vase life increase for ‘Marian Seefruth’
flower was achieved for both dipped (38 days) and
the full spray treatment (39 days) and was signifi-
cantly different from the untreated flower vase life
of 28 days. When only the top of the spathe was
sprayed, the increase was not significantly differ-
ent from full spray but it was sometimes more
variable. Similar results to the different applica-
tion methods were obtained with the anthurium
cultivars ‘Purple Arc’, ‘Blush Oishi’ and ‘Leilani’.
Both dipping the whole flower and spraying were
regarded as equally effective application methods
and the full spray treatment was used in all subse-
quent tests.
Anthuriums were routinely packed in a carton
for 2 days to simulate commercial handling.
When anthuriums were packed and held in the
carton for up to 12 days at 22°C, there was only
a slight decline in vase life after 4 days packed in
the carton (Fig. 2). A significant decline in the
vase life of ‘Ozaki’ anthurium was found when
packed for more than 4 days (Paull, 1984, Unpub-
lished). Anthurium packed in a shipping carton
and held at 22°C had a longer vase life than
flowers held at 8°C. This confirmed an earlier
finding that anthuriums held at 8°C had a shorter
vase life than those held at 14°C or above (Paull,
1987). The BA spray treatment applied to flowers
that had been harvested 3 days before packing,
and, harvested on the day of packing, with BA
applied immediately before or applied after 2 day
 R.E. Paull, T. Chantrachit / Posthar6est Biology and Technology 21 (2001) 303–310
Fig. 2. The interaction between BA (200 mg L − 1 spray) and
time packed in the carton for various lengths of time, on
anthurium (cv. Leilani) vase life. Analysis of variance: BA
application, P =0.00001; BA 5 Storage temperature, P.=
0.05; BA × Storage time, P= 0.01; BA × Storage tempera-
ture× Storage time, P= 0.001; Storage temperature,
P= 0.0001; Storage time, P= 0.0001; Storage temperature ×
Storage time, P =0.0001.
packing gave a similar increase in vase life (Table
2) to those flowers that were harvested on the day
of packing. Whittaker (1993) and Shirakawa et
al., (1964) applied BA after shipping and unpack-
ing, while our data showed that BA can be ap-
plied either before or after packing and shipping.
Anthurium flower size did not significantly infl-
uence the vase life response of ‘Kalapana’ to BA
Table 2
‘Marian Seefurth’ anthurium were harvested from the same block in a uniform area of a commercial shade house, 3 days before
planned packing and on the same day as packing. Those flowers harvested before and on the same day as packing were treated with
200 mg mL−1 BA. Flowers harvested on the same day as packing were treated with BA immediately upon unpacking after 2 days
in a carton at 22°C
Days before packing Vase life (days from harvest)a
No BA treatment
−3 25 c
0 23 c
After unpacking 23 c
a
Means within and between the columns followed by same letter were not significantly different by Duncan–Waller multiple range
test at P50.05, n = 12.
307
applied as either as a dip or spray. Small an-
thurium had a vase life of 29 days dipped and 31
days sprayed, while large flowers had a vase life of
31 and 30 days, respectively, vs. the significantly
lower vase life of 19 and 25 days, respectively, for
the untreated flowers. ‘Leilani’ anthurium har-
vested with 50% open flowers on the spadix had
1.8 fold (28 vs. 49 days) greater vase life after BA
treatment and those with 100% open spadix flow-
ers, a 1.6 fold increase (31 vs. 48 days). The extent
of spadix flower opening at harvest did not signifi-
cantly affect vase life, while the BA response was
statistically significant. These results indicated
that the flower size or spadix flower opening of
more than 50% did not significantly influence the
response to BA. The vase life and degree of
response to BA varied from harvest to harvest,
and BA always increased the vase life of respon-
sive cultivars. ‘Marian Seefurth’ harvested in Au-
gust showed a 1.5 fold greater vase life to BA
treatment and a 1.8 fold and a 1.9 fold increase in
vase life in September and October, respectively,
following BA treatment (data not shown). Sea-
sonal variation in anthuriums vase life has been
previously reported (Paull et al., 1992) and the
increase in vase life following BA treatment also
varied from harvest to harvest. It is possible that
available cytokinin in the anthurium flower could
vary throughout the year, due to environment
conditions and possibly with flower production
per plant.
Treatment of Heliconia with BA delayed both
bract darkening and bract abscission. There was
Increase in vase life times (×)
BA treated
47 a 1.9
43 b 1.9
46 ab 2.0
308 R.E. Paull, T. Chantrachit / Posthar6est Biology and Technology 21 (2001) 303–310

Table 3 nia ‘Sexy Pink’ (Table 3), and, the vase life was
The effect of dipping in BA (200 mg L−1) on ‘Andromeda’
significantly increased from 7 to 21 days when 200
heliconia vase life and either dipping or spraying on ‘Sexy
Pink’ heliconia vase life. The ‘Andromeda’ leaves on the flower mg L − 1 BA was sprayed onto the flowers and to
stem were evaluated separately. Flowers were packed in a 18 days when the flowers were dipped in BA (Fig.
carton for 2 days and held at 22°C before unpacking, 4–5 cm 3). As for anthuriums, the BA could be applied to
cut from stem and placed in deionized water for vase life Heliconia by either spraying or dipping without a
evaluation
significant difference in vase life.
Vase life (days from harvest)a
Benzyladenine treatment increased (1.3 fold)
the vase life of red ginger inflorescence over the
Leaves Flowers untreated control (Table 4). The presence or re-
moval of the leaves from red ginger inflorescence
‘Andromeda’ heliconia
stems did not influence ginger inflorescence vase
No BA treatment 24 b 13 b
BA treated 32 a 31 a
life response to BA (Table 4). The vase life of the
leaves attached to the red ginger inflorescence was
‘Sexy Pink’ heliconia
increased 1.3 fold by BA treatment and this in-
No treatment 7c
Sprayed with BA 21 a crease in vase life was similar to the results of
Dipped in BA 18 b Whittaker (1993). The increase in vase life of red
ginger inflorescences treated with BA varied sig-
a
Means within a column followed by the same letters were nificantly with different harvest dates from 1.4
not significantly different by Duncan–Waller multiple range
test at P50.05, ‘Andromeda’ n= 12, ‘Sexy Pink’ n = 6.
fold to 1.9 fold (Table 5). The extent of this BA
induced increase in vase life was due in part to the
vase life decline of inflorescences harvested from
23 days when harvested in May to 14 days when
harvested in October. The BA treatment reduced
this harvest related decline in vase life and there-
fore could reflect in part to different in vivo levels
of cytokinin.
Other tropical ornamentals showed varied re-
sponses to a BA dip (200 mg L − 1). Pink ginger
flowers and anthurium leaf showed a vase life
increase of 1.6 fold and 2.2 fold, respectively
(Table 6). The responses, were similar to pub-
lished results, in that BA delays senescence in
foliage (Hinkelton, 1991; Han, 1995; Philosoph-
Hadas et al., 1996). The physiological and molec-
ular basis for cytokinin action is based upon
Fig. 3. Effect of BA (200 mg L − 1 spray) on the bract fall and exogenous cytokinin manipulation modifying
bract darkening of the large heliconia ‘Sexy Pink’. Analysis of senescence related gene expression (Hare and van
variance: Bract darkening; BA application, P= 0.0005; days, Staden, 1997). Gan and Amasino (1995) have
P 50.0001; BA application × days, P= 0.0001. Bract fall; BA
shown that plants with increased cytokinin pro-
application, P=0.0001; days, P=0.0001; BA application ×
days, P= 0.0001. duction delays the senescence associated decline in
cytokinin supply, found during monocarpic senes-
cence (Nooden et al., 1990). The exogenous BA
2.4 fold greater vase life of ‘Andromeda’ Helico- applied in the experiments reported here, may
nia leaves and flowers following BA treatment have provided additional cytokinin to delay senes-
(Table 3) and the longer vase life was associated cence. For anthurium, stem clogging and the sub-
with a delay in bract darkening. Bract abscission sequent decline in water uptake has been shown
was also significantly delayed in the large Helico- to be related to reduced vase life (Paull and Goo,
 R.E. Paull, T. Chantrachit / Posthar6est Biology and Technology 21 (2001) 303–310 309

Table 4
Effect of BA (200 mg L−1) spray on the vase life of red ginger inflorescences with and without leaves. Inflorescences were packed
in a carton for 2 days and held at 22°C before unpacking, 4–5 cm cut from stem and placed in deionized water for vase life
evaluation

Inflorescence vase life (days from harvest)a Leaf vase life (days from harvest)

With leaves Without leaves

No BA treatment 26 b 25 b 23 b
BA treated 32 a 32 a 31 a
Analysis of Variance Probability
Presence of leaves 0.375
BA treatment 0.0001
Leaves×BA treatment 0.583

a
Means followed by the same letter were not significantly different by Duncan–Waller multiple range test at P50.05, n =10.

Table 5
The effect of BA (200 mg L−1) spray on the vase life and the increase in vase life of red ginger inflorescences harvested on three
different dates. Inflorescences were packed in a carton for 2 days and held at 22°C before unpacking, 4–5 cm cut from stem and
placed in deionized water for vase life evaluation

Harvest date Vase life (days from harvest)a Increase vase life times (×)

Control BA

96 May 29 23 b 33 a 1.4
96 July 20 19 c 35 a 1.8
96 October 01 14 d 26 b 1.9
Mean 19 31

a
Means within and between the columns followed by same letters were not significantly different by Duncan-Waller multiple
range test at P50.05, n =12.

Table 6
Effect of a BA (200 mg L−1) spray on vase life of different ornamentals. Flowers were packed in a carton for 2 days and held at
22°C before unpacking, 4–5 cm cut from stem and placed in deionized water for vase life evaluation

Species Vase life (days from harvest)a Increase vase life times (×)

Control BA

Bird of Paradise 14 a 14 a 1.0

Pink Ginger 14 b 23 a 1.6
Lycopodium 40 a 28 b 0.7
Beehive Ginger 20 a 17 a 0.9
Bamboo Orchid 58 a 42 b 0.7
Anthurium Leaf 46 b 74 a 2.2
Uluhe Fern Curls 10 a 9 a 0.9

a
Means in the same row, followed by same letter were not significantly different by Duncan–Waller multiple range test at
P50.05, Bird of Paradise and Pink ginger n = 10, Beehive ginger n = 6, and all foliage n =12.
310 R.E. Paull, T. Chantrachit / Posthar6est Biology and Technology 21 (2001) 303–310
1985), and preliminary results suggest that BA
maintained water uptake. BA response may be
another factor controlling anthurium vase life and
possibly related to the large observed seasonal
variation in vase life (Paull et al., 1992) and
cultivar differences in the rate of flower senes-
cence did not relate to BA responsiveness (Table 1
and Fig. 1). A significant increase in vase life
following BA treatment occurred on most an-
thurium cultivars, a modified leaf, Heliconia, leaf-
like bracts, the red and pink bracts of ginger and
foliage material. There was little or no response to
BA on Bird of Paradise flowers, the foliage bam-
boo orchid, beehive ginger, and the fern Ly-
copodium. Differences between harvests in BA
response have been reported above for anthurium
and red ginger. In all cases, BA treated anthurium
and red ginger had a longer vase life throughout
the year than the untreated controls. The current
results showed that cultivars vary in their ability
to withstand the stress of packing and shipping
and that BA increased the ability to withstand
simulated packing and shipping stress.
Acknowledgements
The authors greatly appreciate the initial work
and discussions related to this project by Dr
Jingwei Dai. This is the College of Tropical Agri-
culture and Human Resources journal series 4510.
This research was funded in part by the USDA-
CSREES Special Grant for Tropical Floriculture
93-34199-8355.
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