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Floral induction in tropical fruit trees: Effects of temperature and water

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Journal of Horticultural Science (1994) 69 (3) 397-415

Floral induction in tropical fruit trees: Effects of

temperature and water supply
By S. CHAIKIATTIYOS,1 C. M. MENZEL2 and T. S. RASMUSSEN3
1
Sisaket Horticultural Research Centre, Sisaket 33000, Thailand
2
Maroochy Horticultural Research Station, Queensland Department of Primary Industries,
P.O. Box 5083, Sunshine Coast Mail Centre, Nambour, Queensland 4560, Australia
3
Agricultural Chemistry, Queensland Department of Primary Industries, Meiers Road,
Indooroopilly, Queensland 4068, Australia

SUMMARY
Floral induction in tropical trees generally follows a check in vegetative growth.
However, it is not easy to identify the environmental factors involved in flowering, which
normally occurs during the dry season when temperatures are also often lower. The
separate and combined effects of temperature and water supply on floral induction were
investigated in 'Hass' avocado (Persea americana), 'Lisbon' lemon (Citrus limon), 'Wai
Chee' litchi (Litchi chinensis) and 'Sensation' mango (Mangifera indica). Low
temperatures (15°/l0°C or 15°/l0°C and 20°/15°C compared with 30°/25°C and 25°/20°C)
generally decreased vegetative growth and induced flowering in well-watered avocado,
litchi and mango. A pre-dawn leaf water potential ('l'L) of -1.7 to -3.5 MPa compared
with -0.4 to -0.7 MPa in control avocado and litchi, and a pre-dawn relative water content
(R.W.C.) of 90-93% compared with 97% or above in control mango plants also reduced·
or eliminated vegetative growth, but did not induce flowering. Low temperatures (15°/
10°C Compared With 20°/l5°C, 25°/20°C Or 30°/25°C) and water Stress (pre-dawn 'JfL Of-2.0
to -3.5 MPa compared with -0.7 to -0.8 MPa in controls) reduced or eliminated
vegetative growth in lemon. In contrast to the response in avocado, litchi and mango,
flowering in lemon was very weak in the absence of water stress at 15°/l0°C or outdoors in
Brisbane in subtropical Australia (Lat. 28°S), and was greatest after a period of water
stress. The number of flowers increased with the severity and duration of water stress
(two, four or eight weeks) and was generally greater after constant rather than with cyclic
water stress. In lemon and litchi, net photosynthesis declined with increasing water stress
reaching zero with a midday 'l'L of -3.5 to -4.0 MPa. This decline in carbon assimilation
appeared to be almost entirely due to stomatal closure. Despite the reduction in midday
C02 assimilation, starch concentration increased during water stress, especially in the
branches, trunk and roots of lemon. Leaf starch was uniformly low. The number of
flowers per tree in lemon was strongly correlated with starch in the branches (r 2=77 % ,
P<0.01) and roots (r2=74%, P<0.001). In litchi, starch was lower than in lemon roots and
was not related to flowering.
In separate experiments to test the interaction between temperature and water supply,
low day/night temperatures (23°/18° and l8°/15°C compared with 29°/25°C) reduced
vegetative growth and induced flowering in avocado, litchi and mango. None of these
species flowered at 29°/25°C or as a result of water stress ('l'L of -1.5 MPa compared with
-0.3 MPa for avocado and -2.0 MPa compared with -0.5 MPa for litchi, and R.W.C. of
90-93 % compared with 95-96 % in mango). In contrast, in lemon, flowering was very weak
( <10 flowers per tree) in the absence of water stress (pre-dmyn 'l'L of-2.0 MPa compared
with -0.5 MPa) and was only heavy (>35 flowers per tree) after stressed trees were
rewatered. There were slightly more flowers at 18°/15°C than at 23°/18° and 29°/25°C in
control plants, but no effect of temperature in stressed plants. Starch concentration in the
roots of avocado, lemon, litchi and mango was generally higher at l8°/l5°C and 23°/l8°C
than at 29°/25°C. Water stress increased the starch concentration in the roots of lemon and
398 Floral induction in t~opical crops

litchi and decreased it in avocado. There was no effect in mango. There was a weak
relation (r2=57%, P<0.05) between the number of flowers per tree in lemon and the
concentration of starch in the roots. In contrast, there was no significant relationship
between flowering and starch levels under the various temperature and water regimes in
the other species. In another experiment, only vegetative growth in litchi and mango
occurred at 30°/25°C and only flowering at 15°/10°C. Six weeks of water stress (pre-dawn
'llL of -2.5 MPa compared with -1.0 MPa or higher in litchi, and R.W.C. of 90-93%
compared with 95 % or higher in mango) in a heated glasshouse (30°C days/20°C night
minimum) before these temperature treatments did not induce flowering.
Temperatures below 25°C for avocado and below 20°C for litchi and mango are
essential for flowering and cannot be replaced by water stress. The control of flowering in
lemon over the range of day temperatures from l8°C to 30°C differed from that of the
other species in being mainly determined by water stress. Flowering was generally weak
in well-watered plants even with days at 18°C. Starch did not appear to control flowering.

IN MOST tropical and subtropical fruit trees,
flower initiation usually follows a check in
vegetative growth which occurs during a period
of cold weather or reduced water supply. As
temperatures are normally lower during the dry
season in the tropics and especially in the
subtropics, it is not immediately apparent
whether floral initiation is induced by low
temperature or water stress. Sweet orange
cultivars such as 'Washington Navel', 'Late
Valencia' and 'Frost Valencia' have been
reported to flower with day or night
temperatures below 20°C (Lenz, 1969; Moss,
1976; Hall et al., 1977; Lovatt et al., 1988a). No
information is available on the flowering of
lemon under different temperature regimes,
although 'Lisbon' lemon flowered profusely
after a period of water stress (Nir et al., 1972;
Lovatt et al., 1988b ). 'Tahiti' lime appears to
respond to both low temperatures and drought
(Southwick and Davenport, 1986). Low
temperatures also favoured flower induction in
avocado (Buttrose and Alexander, 1978),
mango (Whiley et al., 1989) and litchi (Menzel
and Simpson, 1988; Menzel et al., 1989), but
there is no clear evidence for a separate and
direct effect of water stress in these species. It is
widely believed that water stress during the dry
season is the factor which induces flowering in
mango in the tropics where the minimum
temperature never falls below 20°C. Little
information is available on the water status of
trees in these locations. In work previously
reported, plants have been grown under a range
of experimental conditions which makes
comparisons between different species difficult.
The trees in our experiments were grown under
the same standard conditions.
Starch reserves have been proposed as
important factors controlling the productivity
of fruit trees (Monselise and Goldschmidt,
1982), although previous reports on the
relationship between flowering and starch
content in tropical tree crops appear to be
inconsistent. The most detailed research has
been conducted in Citrus. Goldschmidt and
Golomb (1982) and Goldschmidt et al. (1985)
found a strong relationship between flowering
and starch associated with cincturing or fruit
removal. However, other workers failed to
show any association between these two factors
(Lewis et al., 1964; Jones et al., 1974). The lack
of response in these studies could be due to the
choice of leaves for sampling, which may not
necessarily reflect the carbohydrate status of
the tree. The role of starch reserves for avocado
flowering is not supported by the observation
that starch concentrations were at their lowest
just prior to floral initiation (Scholefield et al.,
1985). Nevin and Lovatt (1987) also showed
that starch concentration in avocado did not
change with temperature or water stress, but
they did not present any growth or flowering
data. In litchi, there was a good relationship
between flowering and starch concentration
under various temperature regimes, but no
relationship with changes in water supply
(Nakata and Watanabe, 1966; Nakata and
Suehisa, 1969; Menzel et al., 1989).
Suryanarayana (1978) found the highest starch
 S. CHAIKIAITIYos, C. M. MENZEL and T. S. RASMUSSEN
in leaves and stems of several mango
cultivars during flower bud formation
especially during a good flowering year. In
contrast, Patil et al. (1988) reported no
difference in starch at different stages of flower
bud differentiation in 'Alphonso' mango trees,
while Whiley et al. (1989) studying several
mango cultivars, also found no link between
starch levels and flower initiation under
different terp.peratures.
We report the effects of temperature and
water stress on vegetative growth and floral
induction of avocado, lemon, litchi and mango.
The four species were first grown with a
plentiful water supply under four temperature
regimes or under standard temperature
conditions under variable water supply. A
second set of experiments examined the
interaction of temperatures and water stress.
The relationship between flowering and starch
levels was also assessed.
MATERIALS AND METHODS
Temperature experiment
Air-layers of 'Wai Chee' litchi, grafted 'Hass'
avocado, 'Lisbon' lemon, and 'Sensation'
mango trees were grown in ten litre pots of
standard UC. mix in a heated glasshouse. After
·15 months, the trees were transferred to nat-
urally-lit controlled temperature cabinets with
day/night cycles of 15°/10°, 20°/15°, 25°/20° or
30°/25°C from February to July. All plants were
vegetatively dormant at the time of transfer.
The duration of day temperature was 10 h and
mean irradiance about 9.5 MJ m-2 day-1• Vapour
pressure deficit (VPD) during the day was
about 0.9 kPa. Plants in this and all other
experiments were about 0.5 to 1.5 m in height.
When harvested, there was no evidence that
they were pot-bound. This was supported by
the strong shoot growth recorded in some of the
treatments.
A record was kept of daj:es of floral emer-
gence, 50% anthesis and of first appearance and
maturation of vegetative flushes. The propor-
.tions of terminal shoots flowering and dry
weight of the panicles were recorded at 50%
an thesis. Plan~s which had not flowered after 20
weeks were returned to a glasshouse at a day
temperature of 20°C and a minimum night tem-
perature of 10°C for 4-6 weeks. Data are the
399
means for six plants per treatment. Leaf water
potential ('VL) in avocado, litchi and lemon was
measured before dawn and at midday using a
pressure chamber (Ritchie and Hinckley,
1975). Relative water content (R.W.C.) of
mango leaves was determined at the same time
(Turner, 1981), because the latex in mango
stems does not allow accurate measurement of
'VL with the pressure chamber. Data are the
means of six leaves on two successive days.
Relative water content is defined as:
R WC = (fresh wt.-dry wt.) x 100
· · · (turgid wt.-dry wt.)
Turgid weight is determined by soaking the
tissue in water to hydrate fully, and overcomes
the error associated with seasonal and diurnal
changes in dry weight which make comparisons
of water content on a dry weight basis unsatis-
factory (Turner, 1981).
Water stress experiment
Two year old air-layers of 'Wai Chee' litchi
and grafted trees of 'Hass' avocado and 'Lis-
bon' lemon were grown in 14 litre pots of stan-
dard UC. mix and kept in the open at Brisbane
in subtropical Australia (Lat. 28°S) from June
to October. 'Sensation' mango was originally
included, but had to be omitted because of pre-
mature flowering. During the experiment,
photoperiod was 10 to 12 hand irradiance 11.6
to 28.8 MJ m-2 day-1• Mean monthly relative
humidity ranged from 50 to 90% and maximum
and minimum temperatures from 16.6° to
26.8°C and 7.3° to 15.9°C, respectively.
Three treatments were used: control, con-
stant water stress and cyclical water stress. The
pots were enclosed in plastic bags with only the
stem and leaves exposed. There was a gap
around the stem to allow sufficient gas
exchange with the roots. Control plants were
watered daily by weight to field capacity. For
constant water stress, water was added daily as
required to maintain a pre-dawn 'VL of -2.0
MPa, while cyclical stress was established by
drying each tree until the pre-dawn 'VL reached
-2.0 MPa and then rewatering to field capacity.
The interval between cycles for the cyclic stress
treatment varied from two to three weeeks
depending on tree size and atmospheric
demand. Leaf water potential was measured
two or three times a week at the beginning of
400 Floral induction in tropical crops
the experiment and once a week after trees had
reached the required water status. After 12
weeks, all stressed plants were rewatered to
field capacity. Two weeks after the commence-
ment of the experiment, predawn 'l'L from all
trees including controls was measured. Data
are the means for six leaves on two successive
days. The increase in stem extension per branch
was measured just before rewatering and time
of floral emergence noted. In avocado and
litchi, the type, length and weight of the pan-
icles were determined at 50% anthesis, while
for lemon a record was made of the number of
flowers per tree. Data are means for ten plants
per treatment.
Interaction between severity and duration of
water stress
One year old grafted 'Lisbon' lemon and two
year old air-layers of 'Wai-Chee' litchi were
grown in 14 litre pots of oxisol soil, peat and
. sand (2:1:1, pH 6.6) in a standard glasshouse as
described above. During the experiment from
September to February, the photoperiod was
11to13 h, mean monthly irradiance 11.6-28.5
MJ m-2 day-1 , mean monthly relative humidity
82 % , and maximum temperatures varied from
25° to 34°C and minima from 18° to 24°C. Treat-
ments were applied as soon as new vegetative
growth matured. Trees were maintained at a
pre-dawn 'l'L of -2.0 or -3.5 MPa for two, four
or eight weeks and then rewatered to field
capacity as described above. A set of control
trees with a pre-dawn 'l'L above -1.0 MPa was
also included.
Leaf water potential was measured in
stressed trees two or three times a week before,
and once a week after trees had reached the
desired water status. One week after the experi-
ment had started, pre-dawn and midday 'l'L
were determined on all plants including. con-
trols. Data are the means of six leaves from two
successive days. In litchi, records were made of
the proportion of terminal shoots flowering and
time of panicle emergence, while in lemon the
number of flowers per tree and time of first
visible flower bud were recorded. At the end of
water stress treatments, half the trees from each
treatment were harvested and starch concen-
tration in the leaves, branches ( <1 cm diam-
eter), trunks and whole roots determined as
described by Rasmussen and Henry (1990). For
litchi, only the roots were analysed. Data are
the means of three or six trees. Starch is
normally the main storage carbohydrate in the
above-ground tissues of most woody species,
although a few temperate deciduous fruits such
as apple and cherry accumulate soluble car-
bohydrates (Loescher et al., 1990). In tropical
fruit trees such as avocado, seasonal changes in
starch concentration are much greater (2 to
20%) than those for soluble sugars (1 to 3% ),
suggesting that starch is the major pool of
reserve of carbohydrates (Scholefield et al.,
1985). At the end of the eight-week stress treat-
ments, net photosynthesis (Pn) and stomatal
conductance (gs) were measured at midday
using a portable Li-Cor LI-6200 photosynthesis
meter along with pre-dawn and midday 'l'L·
Duplicate leaves from three trees were sampled
from each treatment. During the photosyn-
thesis measurements, average air temperature
was 31.5°C, VPD 1.6 kPa and photon irradiance
449 µmol quanta m-2 s-1•
Morphological aspects of mango bud
differentiation
Three year old grafted 'Sensation' mango
trees were grown in 14 litre pots of oxisol soil,
peat and sand (2:1:1) in a heated greenhouse.
Vegetatively dormant trees were transferred to
glasshouses at 15°/10°C or 30°/25°C in June with
12 h days. The trees at 30°/25°C were main-
tained at a pre-dawn R.W.C. above 97% (con-
trol) or from 90-93% (stressed) as described
above. Trees at 15°/10°C were watered daily,
but no R.W.C. measurements were taken. The
R.W.C. at 15°/10°C for plants watered daily in
other experiments was generally above 96 % .
Stressed trees were rewatered after four weeks.
During the course of the experiment, trees
received a mean photoperiod of 10.5 h,
irradiance of 8.5 MJ m-2 day-1 and relative
humidity of about 65%. Three terminal buds
were sampled from each treatment weekly for
six weeeks. The small leaves surrounding the
apices were dissected using fine forceps and a 3
mm long section including the apex prepared
and fixed in 70% EA.A. (90% ethanol [70% ],
5% formaldehyde and 5% acetic acid). The
samples were dehydrated in a graded series of
ethanol from 50 to 100 % , each for 30 min and
I
 S. CHAIKIATIIYOS, c. M. MENZEL and T. s. RASMUSSEN
critically point-dried using a Denton apparatus.
They were fixed to aluminium stubs using
double-sided tape and sputter-coated with
platinum (approximately 10 nm thick) using a
Microvac Magnetron sputter coater. Observ-
ations were made with a Philips 505 scanning
electron microscope (SEM).
Temperature x water stress experiment
Three year old air-layers of 'Wai Chee' litchi,
grafted 'Lisbon' lemon, 'Hass' avocado and
'Sensation' mango trees were grown in 10 litre
pots of standard U.C. mix in a heated glass-
house with a day temperature of 30°C falling to
a minimum night temperature of 20°C. Dor-
mant trees were transferred to controlled tem-
perature glasshouses at 18°/15°, 23°/18° or
29°/25°C from March to August. Half the plants
were watered daily (controls). The others were
subjected to constant leaf water stress using a
pre-dawn 'l'L of -1.5 MPa for avocado and -2.0
MPa for litchi and lemon, and a R.W.C. of
90-93 % for mango. Plant water staus was
measured two or three times a week before the
trees had reached the proposed 'l'L or R.W.C.
and then once a week. Pre-dawn and midday
water status were measured 45 and 46 d after
the commencement of the treatments on all
plants including controls and data are the
means for eight leaves.
During the experiment, average photoperiod
was 11.5 h, irradiance 11.5 MJ m-2 day-1 , relative
humidity 70% and vapour pressure deficit
(VPD) during the day 0.9 kPa. After 14 weeks,
half the trees from each treatment were har-
vested while the rest were rewatered and main-
tained at 29°/25°C for a further 11 weeks. The
times of floral emergence and 50 % anthesis,
and the proportion of branches flowering (avo-
cado, litchi and mango) or the number of
flowers per tree (lemon) were recorded. The
increment in stem extension and the time and
number of vegetative flushes during the first 14
weeks were also recorded. Leaves and roots
were analysed for starch concentrations as
described earlier. Data are means for four or
eight trees per treatment.
Transfer experiment
Two year old air-layers of 'Wai Chee' litchi
401
and three year old grafted 'Sensation' mango
trees were grown in 14 litre pots of oxisol soil,
peat and sand (2:1:1, pH 6.6) in a heated glass-
house as described above. Water was withheld
from half the trees until the pre-dawn 'l'L
declined to -2.5 MPa in litchi and R.W.C.
declined to 90-93 % in mango. The water stress
was maintained by replacing evaporative
demand daily. In the control trees, pre-dawn 'l'L
was higher than -1.0 MPa for litchi and R.W.C.
higher than 95 % for mango. After six weeks,
the trees were transferred to controlled tem-
perature glasshouses at 15°/10° or 30°/25°C
(12 h days) and all pots were watered to field
capacity for a further ten weeks.
The experiment was conducted during
February to June with the plants receiving a
daily photoperiod of 10 to 13 h. During water
stress, average irradiance was 16.0 MJ m-2 day-1,
relative humidity 78%, the mean daily maxi-
mum temperature 33°C and the mean mini-
mum 19°C. In the controlled temperature
glasshouses during March to June, average
irradiance was 11.7 MJ m-2 day-1 and relative
humidity 75%. At the end of the temperature
treatments, a record was kept of the increase in
stem length. The times of floral emergence, pro-
portion of terminal shoots flowering and dry
weight of panicles were also recorded. Plants
which had not flowered at 15°/10°C after ten
weeks were transferred to a glasshouse main-
tained at 30°/25°C. At the end of the sixth week
of water stress, fully matured leaves were
sampled for starch concentration as described
earlier. Data are the means of six trees per
treatment.
RESULTS
Avocado
Temperature experiment: Flushing occurred
only at 25°/20°C and 30°/25°C and was earlier
and completed more rapidly at the higher tem-
peratures (Figure 1). Not included in these
records are the vegetative flushes that wer~
produced along with the developing inflore-
scences. Flowering occurred in the 15°/10°C and'
20°/15°C treatments, but in the latter plants
only when they were transferred to the stan-
dard glasshouse (Table I). There was a propor-
tion of terminal branches which remained
dormant at 15°/10°C (64%) and 20°/l5°C
 ~

402 Floral induction in tropical crops

Transfer to
standard
30 °C ..,_..,v M
... glasshouse
...
v M
... ...

_
25 °C ... ...
(a) FE A
20 °C ... ... ...
15 °C ... FE
... ...
A ...

Jan Feb Mar Apr May Jun Jul Aug

v
..,_.,. M
30 °C ... ...
25 °C .,._v... M
... ...
(b) v M
20 °C ... ... ... ...
v..,_..,
15 °C ...
Jan Feb Mar Apr May Jun Jul Aug

30 °C ..,_..,v M
...
v
... ..,_..,
M

25 °C ..,_..,v M
... ...
FE
(c) 20 °C ... ... ...Al ' ...
15 °C ... FE
... ..,_..,
A

Jan Feb Mar Apr May Jun Jul Aug

30 °C .,._v... _.,._.,._.,._
M v M v...
...M ...
.,._.,.v v
.,._..,
M ..,_..,
M
25 °C

(d) 20 °C T••••••••••••••••••~••••~
no shoot growth I
FE
15 °C ... ... ...A--.....
0 30 60 90 120 j1so 180Days
Jon Feb Mar Apr May Jun Jul Aug

FIG.1
Effect of temperature on vegetative flushing and flowering in (a) avocado, (b) lemon, (c)
litchi and ( d) mango. Data are the means of six trees. Maximum standard errors of the
interval or duration of growth for these species were 6.2, 8.0, 6.1and7.9 d, respectively. V ==
vegetative flush emergence. M =vegetative flush maturity. FE= floral emergence. A= 50%
anthesis.

(80% ). The interval between floral emergence affected by temperature. Tree water status did
and 50% anthesis was shorter in the 20°/15°C not vary with temperature (Figur.e 2).
regime (Figure 1). In contrast, the amount of Water stress experiment: Pre-dawn 'l'L for well-
flowering and panicle weight were not strongly watered trees two weeks after the commence-
 S. CHAIKIATTIYOS, C. M. MENZEL and T. S. RASMUSSEN 403

TABLE I
Avocado, litchi and mango
Effect of temperature on vegetative growth and flowering in avocado, litchi and mango. No flowering occurred in lemon. Data are
means for six trees± SE
Percentage of terminal shoots

Leafy Leafless Total Panicle dry

Temperature panicles panicles flowering Vegetative Dormant weight (g/tree)

Avocado
15°/10°C 0 34.7 ± 3.3 34.7 ± 3.3 0 65.3 ± 3.3 1.4 ± 0.1
20°/15°C 0 20.5 ± 8.3 20.5 ± 8.3 0 79.5 ± 8.3 0.8 ± 0.3
25°/20°C 0 0 0 100 0
30°/25°C 0 0 0 100 0
Litchi
15°/10°C 0 72.9 ± 8.3 72.9 ± 8.3 0 27.1 ± 8.3 5.4 ± 1.0
20°/15°C 18.7 ± 7.4 32.1±14.4 50.8 ± 12.4 0 49.2 ± 12.4 2.6 ± 0.6
25°/20°C 0 0 0 100 0
30°/25°C 0 0 0 100 0
Mango
15°/10°C 16.7 ± 16.7 83.3 ± 16.7 100 0 0 7.6 ± 1.2
20°115°C 0 0 0 0 100
25°/20°C 0 0 0 100 0
30°/25°C 0 0 0 100 0

ment of the experiment was -0.4 MPa (SE = Lemon

0.01 MPa). Pre-dawn 'lfL decreased to -2.1 MPa
in the constant stress treatment and to -1.7
MPa in the cyclic stress treatment. Stem exten-
sion was 4.4 ± 0.8, 1.9 ± 0.2 and 0.0 cm in the
control, cyclic stress and constant stress treat-
ments, respectively. Water stress also induced
leaf drop, browning of shoot tips and necrosis of
the leaf tips and margins (data not presented).
Only one tree flowered in the cyclical water
stress treatment and one in the control.
Temperature x water stress experiment: Leaf
water status was not affected by temperature
(Figure 3). Pre-dawn 'lfL in stressed trees were
about 1.0 MPa lower, and midday values about
2.0 MPa lower than in control trees. Vegetative
growth occurred only in well-watered plants at
29°/25°C (Table II). Most of the shoots at 18°/
15°C flowered with the stressed trees flowering
25 to 29 days after rewatering (Table II). Total
flowering (with or without water stress) was less
at 23°/18°C than at 18°/15°C. None of the trees
flowered at 29°/25°C or during water stress. The
concentration of starch in th_e roots was higher
at l8°/15°C and 23°/18°C than at 29°/25°C, and
in well-watered plants compared with those
that were water-stressed (Table II). There was
no significant relationship (P>0.05) betweeen
flowering and starch under the various temper-
ature and water regimes. The situation is com-
plicated by the fact that some of the trees (of all
species) had flowered at the time of sampling
for starch determination while others had not.
Temperature experiment: Trees flushed once at
20°/15°, 25°/20° and 30°/25°C, while plants in the
15°/10°C regime only grew when they were
transferred to the standard glasshouse (Figure
1). Once again, flushing was earlier and the
duration shorter at higher temperatures. None
of the trees flowered. Pre-dawn 'IfL was not
affected by temperature (Figure 2). In contrast,
midday values were 0.5-1.0 MPa lower at
20°/15°, 25°/20° and 30°/25°C than at 15°/10°C.
Water stress experiment: Pre-dawn 'lfL for well-
watered trees two weeks after the commence-
ment of the experiment was -0.7 MPa (SE =
0.04 MPa). Pre-dawn 'lfL decreased to -2.2 MPa
in the constant stress treatment and to -2.0
MPa in the cyclic stress treatment. Vegetative
growth was reduced by cyclical stress and pre-
vented by constant stress, whereas a period of
water stress increased the number of plants
flowering and the number of flowers per tree
compared with the control trees (Table III).
The greatest number of flowers was recorded in
the constant water stress treatment where
flowering occurred 18 d after rewatering.
Flowering in the cyclical stress treatment
occurred a week after that in the control trees.
Interaction between severity and duration of
water stress: Plant water status one week after
water was withdrawn is shown in Figure 4. Leaf
water potentials at noon were about 0.5 to 1.0
404 Floral induction in tropical crops

-0.5

-1.0
. ·-·. ..
·-·
0.0 - - - - - - - , - - - - - . , - - - - - . . , . - - - - - , of water stress increased (Figure 6) with the
greatest number of flowers occurring after eight
weeks at a pre-dawn 'l'L of-3.5 MPa (Table IV).
...'I
Leaf drop in the severe water stress treatment
~ -1.5
Avocado was substantial after two weeks, while in the
-2.0 ,______.._ _ _ ~---~--~-~
moderate treatment it was minimal even after

0.0 l eight weeeks (data not presented). Water stress

strongly increased branch, trunk and root
starch compared with the control (Table IV).

·---
-0.5
Starch generally accumulated over time, but
-1.0
there were only slight differences between the
-1.5 r Lemon
·~ • - • levels of stress. Leaf starch was uniformly low.
-2.0 '-'--'-----~--~---~~ The number of flowers per tree was strongly
correlated with starch in the branches (r2 =77%,
P<0.01) and roots (r2 =74%, P<0.001), but less
0.0
so with that in the trunk (r2=44%, P<0.01). The
·-; -0.5 ! I
=======•------· . concentration of starch in the branches was
·~
0..
.e -1.0 ~ correlated with that in the roots (r2 =74%,
•
~::
..J

~ P<0.001). In contrast, branch and trunk starch

Litchi
f were less well correlated (r2=46%, P<0.05).

Temperature x water stress experiment: Leaf

100
water status was not affected by temperature
9s I (Figure 3). Pre-dawn 'l'L in stressed trees were
96
94
92
·-·---
Mango

15 ° 20 ° 25 °
.
• ----.--:::::::==•====:::::·
30
Day temperature (°C)
FIG. 2
Effect of temperature on pre-dawn (circles) and midday
(squares) leaf water potential ('l'L) in avocado, lemon and
litchi and relative water content (R.W.C.) in mango. Data
are the means of six leaves on two successive days. Maxi-
mum standard error of \j/L is 0.06 MPa and of R.W.C. is 0.9%.
MPa lower than at pre-dawn. Moderate water
stress decreased 'l'L by about 1.5 to 2.5 MPa and
severe water stress by about 2.5 to 3.5 MPa
c0mpared with the controls. At the end of eight
weeks of drying, Pn declined with increasing
water stress to reach zero at a midday 'l'L of
about -3.5 MPa to -4.0 MPa (Figure 5). The
response of P n to 'l'L could satisfactorily be
described by a linear relationship (r2 = 90%,
P<0.001). The decline in Pn appeared to be
almost entirely due to stomata! closure with P n
and gs highly correlated (r2 = 97%, P<0.001).
Flowering occurred three to four weeks after
rewatering (Table IV). No flowering occurred
in the controls or in the stress treatments until
after rewatering. The number of flowers
increased strongly as the severity and duration
°
about 1.5 MPa lower, and midday values 2.0 to
2.5 MP a lower than in control trees. No growth
occurred in the lemon trees maintained at a
pre-dawn 'l'L of about -2.0 MPa (Table V). In
the well-watered trees, stem extension was
greater at 23°/18~ and 29°/25° than at 18°/l5°C.
Flowering was very weak in the absence of
water stress and was only heavy after the
stressed trees were rewatered (Table V). There
were slightly more flowers at 18°/15°C than at
23°/18° and 29°/25°C in control plants, but no
effect of temperature in stressed plants. The
concentration of starch in the leaves and roots
was generally higher at l8°/15°C and 23°/18°C
than at 29°/25°C (Table V). Water stress
increased starch concentration in the roots, and
there was a weak relationship between the
number of flowers per tree (y) and root starch
(x):
y = 3.4 x -27.2 (r2 = 57%, P<0.05)
Litchi
Temperature experiment: Flushing occurred
only at 25°/20°C and 30°/25°C and was earlier
and completed more rapidly at the higher tem-
peratures (Table I, Figure 1). Not included in
this record are the vegetative flushes produced
 S. CHAIKIATIIYOS, C. M. MENZEL and T. S. RASMUSSEN 405

18° 23° 29° 150 23° 29°

'c°
0..
-1
·~:::::::::::::::::
c
0.."'
-1
·---·-·
.. ··················•
:§
-;::
·-----·--·
.
] ·····················•

~p. ~
·------ .-----·
-2 -2

... •···················· ·····················•

p.
...
Q)

"' -3 Avocado <;;; -3 Litchi

"' II:
II:

,_,~ .3"'
•·············· .....• ·····················•
-4 -4

18° 23° 29° 18° 23° 29°

·---·-·
100

'c°
0.. g
-;::
·----.~=
-1
03 $
• g
1
95
·····················•··· • ·······················•················
p.

~
01
II:
-2

-3
·--·---·
Lemon
...
"
"'
II:
"
90 ======·===·
Mango

~ -4
·· ....
·····•·········
....... -· ~
0:

FIG. 3
Pre-dawn and midday leaf water potential (o/L) in avocado, lemon and
litchi, and relative water content (R.W.C.) in mango as affected by temper-
ature and water stress. Day temperature shown with nights S°C cooler.
Data are the means of eight leaves over two days. Maximum standard
errors were 0.06, 0.09 and 0.13 MPa in avocado, lemon and litchi and 0.3 %
in mango. Circles = well watered plants. Squares = stressed plants.
Continuous lines = pre-dawn values. Broken lines = midday values.

along with the developing inflorescences. Trees nor floral growth, but remained dormant at 15°/
flowered at 15°/l0°C and 20°/15°C, with panicles l0°C (27%) and 20°/15°C (49% ). Total flower-
emerging earlier at 15°/l0°C, but taking a ing, proportion of leafless panicles and panicle
longer time to reach anthesis. A proportion of dry weight were greater at 15°/10°C than at 20°I
terminal branches produced neither vegetative 15°C. Pre-dawn 'l'L was not affected by temper-

TABLE II
Avocado
Effect of temperature and water stress on stem extension and flowering, and starch concentration in leaves and roots of avocado
trees harvested at week 14. Data are the means of four or eight trees with maximum SE
Percentage of terminal shoots
Stem extension flowering Starch concentration(% of d. wt.)
(cm) before Days to floral
Treatment transfer BeforeA AfterB emergence Leaf Root
Control
18°/15°C 0 82.6 0 80 1.0 13.3
23°/18°C 0 0 70.0 112 1.5 13.2
29°/25°C 13.8 0 0 0.7 10.8
Water stress
l8°/1S°C 0 0 71.4 127 0.8 10.7
23°/18°C 0 0 55.5 122 0.9 10.5
29°/2S°C 0 0 0 0.9 7.0
Temperature means
l8°/15°C 0 41.3 35.7 0.9 12.0
23°/18°C 0 0 62.8 1.2 11.8
29°/25°C 6.9 0 0 0.8 8.8
Water stress means
Control 4.6 27.5 23.3 1.1 12.4
Water stress 0 0 42.3 0.8 9.4
Max. SE 4.9 14.2 24.0 3 0.3 2.7
ABefore transfer to glasshouse at 29°/25°C and rewatering
BAfter transfer to glasshouse at 29°/25°C and rewatering
 i~

406 Floral induction in tropical crops

TABLE Ill
Lemon
Effect of water stress on stem extension and flowering in lemon. Data are means for ten trees ± SE where applicable
Days to first visible
Treatment Stem extension (cm)A No. of plants flowering No. of flowers per tree flower bud
Control 17.3 ± 2.1 4 8.6 ± 4.4 57 ± 2
Cyclical stress 9.2 ± 0.5 9 48.0 ± 14.5 65 ± 2
Constant stress 0 9 62.4 ± 13.4B 84 ±le

AMeasured at the end of stress

B Measured after rewatering
c 18 d after rewatering

ature, while midday values were about 0.5 MPa
lower at 30°/25°C than at 15°/10°, 20°/15° and
25°/20°C (Figure 2).
Water stress experiment: Pre-dawn 'l'L for con-
trol, constant and cyclic stress treatments two
weeks after the commencement of the experi-
ment were -0.48 MPa (SE=0.04 MPa), -2.1
MPa and-1.8 MPa, respectively. No vegetative
growth occurred in the stress treatments and
only a negligible amount in the controls (Table
VI). Almost all the litchis flowered, with trees
in the constant water stress treatment flowering
only after relief of stress (Table VI). Cyclical
water stress also delayed the appearance of the
-1
flower panicles compared with control trees.
Water stress did not affect the type of growth
produced, although constant stress reduced the
weight and especially the length of panicles.
Most panicles were leafless.
Interaction between severity and duration of
water stress: Leaf water potentials one week
after water was withdrawn were about 0.5 to 1.0
MPa lower at noon than at pre dawn (Figure 4 ).
Moderate water stress decreased 'l'L by about
1.5 to 2.5 MPa and severe water stress by about
2.5 to 3.5 MPa compared with the controls. Net
Lempn Litchi
i tll
<;> 0.14
8 €J
0.12 0
1"' 0.10
0

~ 0
0
'-;tj -2 -t
.g
0.08
p_.
§ 0.06
6 -3
<i
11 0.04
~ -4
e
+-'
q ~ 0.02
Q)
Lemon
+-'
0 I
0.. ';' 12
H
+-'
Q)

ro
-1
0
u
"' 10 s 0
~ -2
......
roQ)
.....:i -3

4
-4

Litchi
018
Control Moderate Severe -4 -3 -2 -1 -4 -3 -2 -1

Water stress treatment Midday leaf water potential

FIG. 4
Pre-dawn and midday leaf water potential ('l'L) of lemon and
litchi in the different water stress treatments of the severity
x duration water stress experiment. Data are the means of
six leaves on two successive days recorded one week after
water was withdrawn. Clear bars= pre-dawn 'l'L· Solid bars=
midday 'l'L· Vertical bars indicate half standard errors.
(MP a)
FIG. 5
Relationship between stomatal conductance (g,) and net
photosynthesis (Pn) with leaf water potential ('l'L) in lemon
and litchi in the severity x duration water stress experiment.
Data are the means of two leaves per tree recorded at mid-
day at the end of water stress.
 S. CHAIKIAITIYOS, C. M. MENZEL and T. S. RASMUSSEN 407

IV
TABLE
Lemon
Effect of water stress on flowering and starch concentration in lemon. Data are means for three trees with maximum SE
Days from
relief of stress Starch concentration(% of d. wt.l
to first visible Number of
Treatment flower bud flowers per tree Leaves Branches Trunk Roots
Control 0 0.3 1.2 2.6 4.6
Two weeks of stress
Moderate 24 7.3 0.7 2.2 4.7 11.7
Severe 22 17.0 0.4 2.0 4.8 9.5
Four weeks of stress
Moderate 27 33.0 0.6 2.3 4.8 12.9
Severe 22 45.3 0.7 2.9 4.9 14.8
Eight weeks of stress
Moderate 25 50.0 0.3 4.9 5.7 18.5
Severe 20 75.3 0.7 5.9 5.8 18.9
Stress means
Moderate 27 30.1 0.5 3.1 5.1 14.4
Severe 22 45.8 0.6 3.6 5.2 14.4
Duration means
1\vo weeks 26 12.2 0.5 2.1 4.8 10.6
Four weeks 25 39.2 0.6 2.6 4.9 13.9
Eight weeks 22 62.7 0.5 5.4 5.8 18.7
Max. SE 3 8.6 0.3 0.6 0.7 3.8
A At the end of water stress

photosynthesis (Pn) declined with increasing P<0.001), with Pn and gs also highly correlated
water stress to reach zero at a midday 'l'L of (r2=88%, P<0.001). Average Pn and gs for litchi
about -4.0 MPa (Figure 5). Once again, the were about half those for lemon. No growth
response of P n to 'l'L could satisfactorily be occurred during the period of moisture stress
described by a linear relationship (r 2=86 % , and dormant buds grew out as vegetative
shoots once the trees were rewatered (Table
100
VII). None of the trees :flowered. Vegetative
Q) growth did not recover to control rates after
Q)
H
......, rewatering. Leaf drop in the moderately
80 stressed trees was very slight, but many leaves
H 0
Q)
p... 0 were lost with two weeks of severe stress (data
rn
H 60 0 not presented). Starch accumulated slightly in
Q)
~ 0 the roots with the duration of water stress
,S
'+-< 40
00 (Table VII). Starch levels were lower than in
0
'+-< 0 lemon roots and were not related (P>0.05) to
0
H
flowering.
Q) 20
..0 0
s
;:J §
0 Temperature x water stress experiment: Leaf
z 0 water potential was about 0.5 to 1.0 MPa lower
at 29°/25°C than at 18°/15°C and 23°/18°C in
0 5 10 15 20 25 30 control plants at midday (Figure 3). Pre-dawn
Cumulative stress (MPa x weeks)
'l'L in stressed trees were about 1.5 MPa lower,
and midday values 2.0 to 3.0 MPa lower than in
FIG. 6 control trees. Vegetative growth occurred only
Relationship between the number of flowers per tree (y) and
cumulative stress (x) in lemon during the severity x duration in well-watered plants at 29°/25°C (Table VIII).
water stress experiment. Data are for stressed trees from Flowering occurred only at 18°/15°C and was
single trees (open circles). A zero stress is assumed for the delayed by water stress. Water stress increased
control plants (closed circle). Regression is y = 1.8 + 2.8 x
(r2 = 92% P<0.001). Leaf water potential measurements starch concentration in the roots (Table VIII).
have been converted to positive integers. Low temperatures (18°/15° and 23°/18°C vs. 29°/
408 Floral induction in tropical crops

V
TABLE
Lemon
Effect of temperature and water stress on stem extension and flowering, and starch concentration in leaves and roots of lemon
trees harvested at week 14. Data are means of four or eight trees with maximum SE
Starch concentration
Stem extension No. of flowers (%of d. wt.)
before transfer - - - - - - - - - - - Days to floral - - - - - - - - - - -
Treatment (cm) BeforeA AfterB emergence Leaf Root
Control
18°/l5°C 18.2 8.0 0 96 7.0 12.4
23°/l8°C 25.8 2.2 0 85 4.5 9.7
29°/25°C 24.5 0 0 1.4 7.8
Water stress
18°/15°C 0 0 36.2 120 5.5 20.5
23°/l8°C 0 0 38.5 121 1.2 18.5
29°/25°C 0 0 37.7 123 0.9 13.9
Temperature means
l8°/15°C 9.1 4.0 18.1 6.3 16.4
23°/l8C 12.9 1.1 19.3 2.8 14.1
29°/25°C 12.3 0 18.9 1.1 10.8
Water stress means
Control 22.8 3.4 0 4.3 10.0
Water stress 0 0 37.5 2.5 17.6
Max. SE 9.1 4.0 18.4 3 2.9 1.8
ABefore transfer to glasshouse at 29°/25°C and rewatering
B After transfer to glasshouse at 29°/25°C and rewatering

25°C) increased starch concentration in the at 25°/20°C and 30°/25°C (Figure 1). Flushing
roots, but only in control plants. There was no was earlier and completed more rapidly at the
significant relationship (P>0.05) between higher temperatures. Flowering only occurred
flowering and starch under the various temper- at 15°/l0°C and most of the panicles were leaf-
ature and water regimes. less (Table I). At 20°/15°C, the branches
remained dormant producing neither vegeta-
Transfer experiment: Vegetative growth tive nor floral growth. Tree water status varied
occurred in the 30°/25°C regime and flowering only slightly with temperature (Figure 2).
in the 15°/l0°C regime (Table IX). The previous
water stress reduced stem extension during this Morphological aspects of bud differentiation:
period compared with the rate in control trees Reproductive growth was first visible in the ter-
(13.7 ± 1.5 cm vs. 20.4 ± 1.0 cm), but had no minal buds of well-watered trees grown at 15°/
effect on flowering or panicle weight. In con- 10°C after five weeks. The buds became swollen
trast, water stress increased leaf starch com- and floral primordia which would have devel-
pared with control trees (Table IX). oped into primary panicle branches were
formed in the axils of the leaves, indicating an
Mango early stage of flower bud differentiation
Temperature experiment: The trees only flushed (Figures 7 and 8). In contrast, terminal buds of
TABLE VI
Litchi
Effect of water stress on stem extension and flowering of litchi. Data are means of ten trees ± SE where applicable
Percentage of terminal shoots
Stem No. of Days to Panicle Panicle
extension plants floral Leafy Leafless Total d.wt. length
Treatment (cmt flowering emergence panicles panicles flowering Vegetative (g/tree) (cm)
Control 1.3 ± 0.5 10 60 4 2.9 ± 2.1 80.5 ± 7.4 83.4 ± 5.8 16.6 ± 5.8 2.5 ± 0.2 9.0 ± 0.4
Cyclic stress 0 10 75 3 0 97.8 ± 1.5 97.8 ± 1.5 2.2 ± 1.5 2.3 ± 0.3 7.8 ± 0.3
Constant stress 0 9B 114 le 5.3 ± 2.9 71.6 ± 9.2 76.9 ± 9.8 23.l ± 9.8 1.7 ± 0.2 2.5 ± 0.4
A Measured at the end of stress
BFlowered after rewatering
c Thirty days after rewatering
 S. CHAIKIATIIYos, C. M. MENZEL and T. S. RASMUSSEN 409

VII
TABLE
Litchi
Effect of water stress on shoot growth and starch concentration in roots of litchi. No trees flowered. Shoots which were not
vegetative were dormant. Data are means for three trees with maximum SE
Bud break (days from Percentage of shoots Starch concentration in
Treatment relief of stress vegetative roots ( % of d. wt. )B

Control 100 1.0

Two weeks of stress
Moderate 35 91.5 2.2
Severe 29 76.1 1.4
Four weeks of stress
Moderate 30 90.6 3.2
Severe 21 80.0 2.6
Eight weeks of stress
Moderate 29 60.6 4.4
Severe 34 67.5 2.7
Stress means
Moderate 80.9 3.3
Severe 74.5 2.2
Duration means
Two weeks 83.8 1.8
Four weeks 85.3 2.9
Eight weeks 64.1 3.6
Max. SE 3 20.4 1.2
AFrom the beginning of the experiment
B At the end of water stress

well-watered plants at 30°/25°C remained veg- Temperature x water stress experiment: Relative
etative with no signs of panicle initiation after water content was not affected by temperature,
five weeks (Figure 9). Terminal buds of trees but was about 4 % lower in stressed trees than in
with a R.W.C. of 90-93% remained dormant controls (Figure 3). Flushing only occurred in
during the four weeks of water stress (Figure well-watered plants at 29°/25°C (Table X). In
10) and grew out as vegetative shoots 1-2 weeks contrast, flowering occurred only at 18°/15°C
after rewatering (Figures 11 and 12). and was delayed by water stress. As with the
TABLE VIII
Litchi
Effect of temperature and water stress on stem extension and flowering, and starch concentration in leaves and roots of litchi trees
harvested at week 14. Data are means of four or eight trees with maximum SE
Percentage of terminal shoots Starch concentration
Stem extension flowering (%of d. wt.)
(cm) before Days to floral - - - - - - - - - - -
Treatment transfer BeforeA AfterB emergence Leaf Root
Control
18°/15°C 0 88.0 0 69 1.7 3.3
23°/l8°C 0 0 0 0.5 2.8
29°/25°C 11.9 0 0 0.3 1.5
Water stress
18°/15°C 0 0 58.7 127 1.7 4.3
23°/l8°C 0 0 0 1.3 4.8
29°/25°C 0 0 0 1.1 5.2
Temperature means
18°/15°C 0 44.0 29.4 1.7 3.8
23°/l8°C 0 0 0 0.9 3.8
29°/25°C 6.0 0 0 0.7 3.3
Water stress means
Control 4.0 29.4 0 0.8 2.5
Water stress 0 0 19.7 1.3 4.8
Max. SE 4.2 11.9 21.3 3 0.5 0.8
ABefore transfer to glasshouse at 29°/25°C and rewatering
B After transfer to glasshouse at 29°/25°C and rewatering
410 Floral induction in tropical crops
TABLE IX
Litchi and mango
Effect ofwater stress on flowering and leafstarch concentration in litchi and mango. Trees grown at 15°/l 0°C and transferred to 30°!
25°C after ten weeks. All trees remained vegetative when maintained continuously at 30°/25°C. Data are means for six trees ± SE
Percentage of
terminal shoots
Treatment flowering
Litchi
Control 100
Previous water stress 100
Mango
Control 100
Previous water stress 100
A After ten weeks
other species, starch concentration was sub-
stantially higher in roots than in leaves (Table
X). The average concentration of root starch
was similar in mango, avocado and lemon (7 .9
to 13.8% ), but much lower in litchi (3.7% ). The
concentration of starch in the roots was higher
at 18°/15°C and 23°/l8°C than at 29°/25°C, but
water stress had no effect. There was no signifi-
cant relationship (P>0.05) between flowering
and starch.
Transfer experiment: Trees grew vegetatively in
the 30°/25°C regime and flowered in the 15°/
10°C regime (Table IX). The previous water
stress had no effect on subsequent vegetative
growth (20.2 ± 1.5 cm in control vs. 23.3 ± 1.5
cm in stressed plants) or on flowering or panicle
weight, but increased leaf starch slightly (Table
IX).
DISCUSSION
Vegetative growth was reduced or prevented
in the tropical fruit trees by low temperatures
and water stress. In contrast, the four species
behaved quite differently in their flowering
response. Low temperatures, but not water
stress, induced flowering in avocado, litchi and
mango. In lemon, flowering was weak in the
absence of water stress and was not influenced
greatly by changes in temperature.
Avocado, litchi and mango
Low temperatures favoured flower initiation
· in avocado, litchi and mango, but delayed
flower development. The promotion of flower-
ing at low temperature was not associated with
water stress as determined by pre-dawn and
midday 'l'L or R.W.C. The incomplete flowering
of litchi at 15°/l0°C in our study compared with
Floral emergence Panicle dry Leaf starch
(days from relief weight concentration
of water stress t (g/tree) (%of d. wt.)
79 ± 1 12.2 ± 2.1 1.0 ± 0.1
76 ± 2 15.7 ± 3.2 1.9 ± 0.3
76 ± 1 39.7 ± 3.9 1.1 ± 0.1
83 ± 1 34.8 ± 3.4 1.8 ± 0.2
reports by Menzel and Simpson (1988) and
Menzel et al. (1989) possibly reflects the imma-
turity of some of the shoots at the start of the
experiment.
Flowering in mango occurred only at 15°I
10°C. The pattern of flower differentiation was
similar to that reported by Moncur (1988), but
our study showed an earlier stage of differ-
entiation. Since floral differentiation was first
visible after five weeks, this approximates to the
critical 'chilling' period for this cultivar,
although, the critical temperature and duration
may vary with cultivar. Nunez-Elisea and
Davenport (1991) reported that mature, dor-
mant stems of 'Tommy Atkins' required 30 to
40 d of chilling. In the study by Whiley et al.
(1989) with cultivars from both tropical and
subtropical climates, only 'Florigon' flowered
at 20°/15°C while eight of the remaining nine
flowered at 15°/10°C. The duration of the treat-
ment (20 weeks) was much longer than that
used in this study and the exact time of panicle
emergence was not indicated.
The temperature at which the transition from
vegetative to floral growth occurs varies with
the species, but generally lies between 15° and
25°C in the subtropical avocado, litchi and
mango. In 'Hass' avocado, 23°/l8°Cwas close to
the critical point, since flowering in this treat-
ment was later and weaker than at 18°/l5°C.
There was a neutral temperature range of 23°/
l8°C in litchi and mango where neither flower-
ing nor vegetative growth occurred. It is poss-
ible that a similar range between 23°/18°C and
29°/25°C exists in avocado. Suppression of veg-
etative growth thus will not induce flowering, at
least in litchi and mango. The response totem-
perature is unique and not replaced by water
stress.
 S. CHAIKIATIIYOS, C. M. MENZEL and T. S. RASMUSSEN 411

FIGS. 7-12
Scanning electron micrograph of the terminal apex in mango under various temperature and water regimes, (left to right from above).
7. After five weeks at 15°/10°C showing an early stage of floral development.
8. Same as Fig. 7 but showing developing primary panicle branch.
9. After five weeks at 30°/25°C showing vegetative stage and absence of floral development.
10. After four weeks at 30°/25°C and a relative water content (R.W.C.) of 90-93 % showing dormant stage with pre-existing leaf
primordia and no signs of floral development.
11. Same as Fig. 10 but after one week of rewatering showing young lateral vegetative buds in the axils of existing leaves. No
new leaves or inflorescences.
12. Same as Fig. 10 but two weeks after rewatering showing lateral buds in the axils of existing leaves. L =dissected leaf base. P
= primary panicle branch primordium. A = apical meristem of primary panicle branch. F = floral primordium. B = bud
scale. T =terminal meristem. Ax axillary meristem.
412 Floral induction in tropical crops

TABLEX
Mango
Effect of temperature and water stress on stem extension and flowering, and starch concentration in leaves and roots of mango
trees harvested at week 14. Data are means of four or eight trees with maximum SE
Percentage of terminal shoots Starch concentration
Stem extension :flowering (%of d. wt.)
(cm) before Days to floral
Treatment transfer BeforeA After13 emergence Leaf Root
Control
l8°/15C 0 25.3 0 95 1.4 10.5
23°/l8°C 0 0 0 0.9 8.1
29°/25°C 19.7 0 0 0.8 5.4
Water stress
l8°/15°C 0 0 31.6 131 1.0 8.5
23°/l8°C 0 0 0 1.3 8.4
29°/25°C 0 0 0 0.8 6.5
Temperature means
l8°/15°C 0 12.7 15.8 1.2 9.5
23°/18°C 0 0 0 1.1 8.2
29°/25°C 9.9 0 0 0.8 5.9
Water stress means
Control 6.6 18.4 0 1.0 8.0
Water stress 0 0 10.6 1.0 7.8
Max. SE 7.0 12.4 18.3 3 0.2 1.2
A Before transfer to glasshouse at 29°/25°C and rewatering
8
After transfer to glasshouse at 29°/25°C and rewatering

Water stress prevented vegetative growth in
avocado, litchi and mango and delayed flower-
ing until the trees were rewatered. Our data do
not support the common belief that drought
induces flowering in certain tropical fruit trees,
especially mango. They do, however, confirm
the results for litchi reported by Menzel et al.
(1989), who found that trees which had been
subjected to water stress flowered only after
they were transferred to a glasshouse at 15°I
10°C. They suggested that water stress can pro-
mote flowering in litchi through inhibition of
vegetative growth, but is not essential. As the
trees in the present experiment were appar-
ently fully dormant, no further effect of water
stress would be expected.
The role of water stress in tropical mango
flowering is yet to be resolved. 'Sensation' is a
Florida selection and would be expected to
have a lower 'chilling' temperature for flower-
ing than more tropical cultivars. The behaviour
of young trees with limited root development
may be quite different from that of mature trees
in the field. Wittwer (1991) examined the
effects of irrigation on the productivity of 'Fas-
cell' mango at Nelspruit in subtropical South
Africa (Lat. 25°S) over three years and found
no significant difference in average yield
between trees irrigated all year and those
where irrigation was withdrawn from May to
July. The winter irrigation appears to have ben-
efited spring flowering and not vegetative
growth, suggesting that a dry period before
May might have been more appropriate as
flowers appear in June. Average minima at Nel-
spruit between May and July are <l0°C. Irri-
gation experiments are needed to evaluate the
role of water stress on mango flowering under
more tropical conditions.
Lemon
Our results indicate that there must be a dif-
ferent mechanism for control of flowering in
lemon. Vegetative growth was absent or poor at
15°/10°C or l8°/lS°C and was enhanced pro-
gressively at higher temperature. Although a
few flowers were produced in the controls at
18°/15°C and 23°/l8°C, flowering was much
greater after water stress. The failure of lemon
trees to flower at 15°/10°C may indicate that this
is a transitional temperature range at which
neither flowering nor vegetative growth occurs.
Tree water status in this study was only slightly
affected by temperature and was not low
enough to indicate plants under water stress.
The number of flowers was related to the
severity and the duration of water stress, with a
tendency to more intense flowering under con-
stant water stress. None of the trees, however,
flowered until rewatered. These responses were
 s. CHAIKIATTIYOS, c. M. MENZEL and T. s. RASMUSSEN 413

associated with a reduction in the amount of
vegetative growth. The maximum flower pro-
duction occurred under conditions which led to
substantial leaf drop. It is apparent that full leaf
cover is not essential for floral induction. How-
ever, since such leaf loss would appear to be
deleterious for fruit production, it is suggested
that less severe water stress may be sufficiently
effective to promote flowering. In Sicily,
:flowering and production of lemon is managed
withholding water during summer until the
trees wilt which corresponds to a pre-dawn 'IfL
of about -1.3 MPa (Barbera et al., 1985). Some-
times the trees may be stressed too severely
causing root damage and poor fruit quality.
Photosynthesis and starch
Net photosynthesis in lemon and litchi
declined with increasing water stress to reach
zero at a midday 'lfL of-3.5 to -4.0 MPa and this
decline appeared to be almost entirely due to
stomata! closure. Evidently, there must have
been some photosynthetic activity in the early
morning or late afternoon in stressed lemon
trees as the accumulation of starch in the roots
of trees with a pre dawn 'lfL of -3.5 MPa did not
appear to have been at the expense of the other
plant parts. Leaf starch was a poor indicator of
the effects of water stress and starch in the rest
of the plant. In litchi, starch concentration was
determined only on root samples, as it was the
most sensitive indicator of water stress in
lemon. Starch concentration was much lower
than in lemon and was only slightly increased
with water stress.
These experiments provide little evidence
for a causal role of starch in the :flowering of
tropical fruit trees. Starch concentration was
generally higher at lower temperatures, but was
higher in water stressed plants only in lemon
and litchi. High starch concentrations were
generally associated with lack of vegetative
growth rather than with :flowering. Our data
suggest that increased starch contents may be
coincidental with :flowering, but not the cause.
Data collected by Lovatt et al. (1988b) demon-
strate the difficulty of relating flowering and
starch levels. They showed that the number of
flowers in Citrus increased in proportion to the
number of weeks at low temperatures (15° -
18°C day/l0°-l3°C night), with no significant
effect of the duration of low temperatures on
leaf starch concentration which was highly vari-
able. However, there was a correlation
(r2=56%, P<0.001) between the number of
flowers per tree and the concentration of starch
in the different treatments one week after the
plants were transferred to a warmer glasshouse
at 24°/19°C.
We would like to thank the staff at the
Botany Department, University of Queensland
for their help during this project, especially Dr
W. G. Slater for his guidance during the experi-
mental work and preparation of the manu-
script. Financial support by the Australian
International Development Assistance Bureau
(AIDAB) for S. Chaikiattiyos is gratefully
acknowledged.

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(Accepted 24 October 1993)

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