Scientia Horticulturae 96 (2002) 67–79

Root temperature effects on ‘Arkin’ carambola

(Averrhoa carambola L.) trees
II. Growth and mineral nutrition
Hilary L. Georgea,b, Frederick S. Daviesb,
Jonathan H. Cranea,*, Bruce Schaffera
a
Tropical Research and Education Center, University of Florida, IFAS, Homestead, FL 33031, USA
b
Horticultural Sciences Department, University of Florida, Gainesville, FL 32611, USA
Accepted 5 June 2002

Abstract

The effects of root temperature on plant growth and leaf nutrient element concentrations of 1-year-
old ‘Arkin’ carambola (Averrhoa carambola L.) trees in containers were investigated. Plants were
grown in controlled root temperature chambers in a glasshouse at root temperatures of 5, 10, 20, 25,
or 38 8C. The rate of leaf chlorosis and abscission was higher for trees at root temperatures of 5 and
10 8C compared to trees at the other root temperatures. Tree height, stem diameter, and length and
number of axillary shoots increased as root temperatures increased. Also, the percentage of chlorotic
and abscised leaves decreased as root temperature increased. Plant fresh and dry weights were
generally greatest at root temperatures of 20 and 25 8C. There were no differences in leaf N, P, and K
concentrations among root temperature treatments. However, leaf Ca, Mg, Fe, and Zn concentrations
were greater for trees at 5 8C than trees at the other root temperatures. Based on these results,
increased growth and yield of carambola trees as a result of mulching in orchards may be due to
increased root-zone soil temperature that has been observed for mulched trees.
# 2002 Elsevier Science B.V. All rights reserved.

Keywords: Star fruit; Chlorosis; Abscission; Plant height

1. Introduction

Temperature is the principle environmental factor that influences the adaptability

of tropical woody species to subtropical climates (Berry and Bjorkman, 1980). Ngah
et al. (1989) reported that the ideal air temperature range for growth of carambola
*
Corresponding author. Tel.: þ1-305-246-7001; fax: þ1-305-246-7003.
E-mail address: jhcr@mail.ifas.ufl.edu (J.H. Crane).

0304-4238/02/$ – see front matter # 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 3 0 4 - 4 2 3 8 ( 0 2 ) 0 0 0 9 1 - 2
68 H.L. George et al. / Scientia Horticulturae 96 (2002) 67–79

(Averrhoa carambola L.) trees is 21–32 8C. In south Florida, temperatures for growing
carambola trees are suitable for 9–10 months of the year. However, periods of low tem-
peratures during December through February often decrease tree growth and alter tree
development (Campbell et al., 1985). In a study of the effect of mulch on 5-year-old carambola
trees in an orchard in south Florida, increasing root temperatures as a result of mulching,
increased tree growth (George et al., 2001a,b). Thus, the decreased rate of tree growth
during the winter months in south Florida may be due to a decrease in root temperatures.
The mechanism by which root temperatures affect tree growth of carambola has not been
elucidated. George et al. (2001c) investigated the effects of mulching on root temperature
and leaf nutrient element concentrations in carambola. Despite increased growth as a result
of higher root temperatures, mulching had no significant consistent affect on leaf nutrient
element content over time (George et al., 2001c).
An understanding of nutrient element uptake as it relates to growth of carambola under
various soil temperatures should provide useful information for management and selection
of planting areas for carambola cultivation. Studying carambola nutritional and growth
responses to root temperature in controlled root temperature chambers in a glasshouse
allow root temperature responses to be quantified and easily separated from responses to
other environmental variables such as ambient air temperatures. The objectives of this
study were to determine the effects of root temperature on plant nutrition, growth, and
development of container-grown carambola trees in controlled conditions.

2. Materials and methods

2.1. Plant material and care

One-year-old ‘Arkin’ carambola trees grafted onto open pollinated ‘Golden Star’
carambola rootstocks were grown in 3.8 l containers in a well-drained artificial medium
(George et al., 2002). The trees were kept in a sunlit glasshouse at day/night temperatures
ranging from 30/25 to 25/20 8C, irrigated to container capacity three times each week, and
fertilized every 4 weeks prior to commencement of the experiments with Osmocote1 Plus
controlled-release and MicrolificTM iron as EDDHA fertilizer as described by George et al.
(2002).

2.2. Treatments

Prior to the experiment, trees ranged in height from 87.6 to 137.8 cm with stem
diameters of 6.8–11.1 mm. Six weeks prior to placing plants in the root chambers, plants
were categorized by size so that each treatment group had plants with similar height means
and standard deviations; trees were then pruned as described by George et al. (2002). Two
weeks before treatments were initiated, trees were transferred to another glasshouse
containing five root temperature chambers. Mean ambient day/night air temperatures in
the glasshouse ranged from 27/16 to 38/25 8C.
Beginning from 3 February 1999, trees were divided into the following root temperature
treatments: 5, 10, 20, 25, and 38  1 8C by placing them in one of five root temperature
 H.L. George et al. / Scientia Horticulturae 96 (2002) 67–79 69

chambers, each set at the specified temperature. Due to limitation of the number of root
chambers (one per temperature treatment), the experiment was replicated three times as
described by George et al. (2002). Thus, root temperature treatment was replicated three
times and each replication consisted of six trees. During regrowth, the terminal axillary
shoot was trained to allow upward extension growth. Plants in the root chambers were
irrigated to container capacity three times a week with water at the same temperature as
the roots.

2.3. Root temperature chambers

The root temperature chambers were thermostatically controlled freezers, modified for
use as root temperature chambers as described by George et al. (2002). The temperature of
the ambient air and soil in the plant containers in each growth chamber was continuously
monitored and recorded using a Hobo1 H8 Pro Series temperature/external temperature
logger (Onset Computer Corporation, Pocasset, MA) and a Fisher1 mercury thermometer
(Fisher Scientific Co., IL).

2.4. Growth measurements

Plant height, stem diameter, axillary shoot growth (length), the number of developed
nodes per axillary shoot, leaf chlorosis, leaf abscission, and fresh and dry weights were
determined for plants in all root temperature treatments. All growth measurements, except
fresh and dry weights, were done at the beginning (day 0) of each replication, and repeated
at 14-day intervals. All growth measurements were made on each of the six trees per
replication in all three replications with the exception of leaf chlorosis and abscission
measurements, which were made on three randomly selected trees per replication.
Tree height was measured on the regrowth of the apical shoot arising from the main stem
after pruning. The stem diameter was measured at a 10 cm height above the graft union
using a micrometer caliper. Axillary shoot growth and the total number of developed nodes
per shoot were determined by measuring shoot extension growth, and counting the number
of nodes present on the three axillary shoots that developed just below the tree apex. The
number of chlorotic and abscised leaves was determined for the 10 lowest leaves on the
main stem and expressed as a percentage. At the end of each replication, fresh and dry
weights of leaves, stems (including axillary branches and shoots), and roots were
determined. Tissue samples were oven-dried to constant weight at 70 8C and the dry
weights recorded.

2.5. Leaf mineral element content

Leaf mineral element concentrations were determined for the most recently matured leaf
(fourth or fifth) on the main stem apex of each tree in each treatment. Leaves were sampled
at the beginning and end of each replication. Leaves were compiled into a composite
sample for each treatment for leaf element analysis. Leaf samples were gently rinsed under
running de-ionized water for about 1 min. Samples were oven-dried at 70 8C for 48 h and
ground to a powder in a cyclone grinder (UDY Corporation, Fort Collins, CO).
70 H.L. George et al. / Scientia Horticulturae 96 (2002) 67–79

Nitrogen concentrations were determined by the total Kjeldahl nitrogen (TKN) method
described by Hanlon et al. (1994). After Kjeldahl digestion of 0.2 g of ground tissue for
each sample, TKN concentrations were quantified with a rapid-flow automated spectro-
photometer (Alpchem Corp., College Station, TX). One gram of ground tissue for each
sample was ashed in a ISOtemp1 Muffle Furnace model 58 (Fisher Scientific Company,
Atlanta, GA), extracted in HCl, and P, K, Mg, Ca, Fe, Mn, Zn, Cu, and B concentrations
were determined with an inductively coupled argon plasma spectrometer (TJA Solutions,
Franklin, MA) as described by Hanlon et al. (1994).

2.6. Statistical analysis

The treatments were arranged as a randomized complete block (RCB) in time with three
replications (times). Data were analyzed using the Statistical Analysis Systems computer
software package (SAS Institute, 1996). Main effects (after separating interacting variables
if necessary) were analyzed over time within each replication using repeated measures
analysis. On the final measurement day (day 42) of each replication, regression analyses

Fig. 1. Effect of root temperature on the (a) increase in plant height, and (b) the increase in stem diameter of 1-
year-old, container-grown ‘Arkin’ carambola trees from 3 February to 8 June 1999. Each data point is the mean
of six trees from each of three replications per treatment. Vertical bars indicate 1 S.E. Measurements were
made at 14-day intervals. The regression lines for (a) height over time are represented by the equations: for 5 8C,
y ¼ 0:59 þ 0:11x, r2 ¼ 0:42; for 10 8C, y ¼ 1:85 þ 0:22x, r2 ¼ 0:42; for 20 8C, y ¼ 0:95 þ 0:43x,
r 2 ¼ 0:74; for 25 8C, y ¼ 1:13 þ 0:75x, r 2 ¼ 0:88; for 38 8C, y ¼ 0:04 þ 0:61x, r2 ¼ 0:91. The regression
lines for (b) stem diameter over time are represented by the equations: for 5 8C, y ¼ 0:01 þ 0:01x, r2 ¼ 0:77;
for 10 8C, y ¼ 0:18 þ 0:02x, r 2 ¼ 0:74; for 20 8C, y ¼ 0:31 þ 0:02x, r2 ¼ 0:56; for 25 8C,
y ¼ 0:07 þ 0:03x, r 2 ¼ 0:82; for 38 8C, y ¼ 0:27 þ 0:04x, r2 ¼ 0:59. For all equations, P  0:01.
 H.L. George et al. / Scientia Horticulturae 96 (2002) 67–79 71

were done with temperature as the independent variable and leaf nutrient concentrations,
growth, and development variables as the dependent variables. Leaf mineral element
concentrations were analyzed by non-linear regression analysis at the beginning and end of
each replication (TableCurve 2D, SPSS Science Software, Chicago, IL).

3. Results

3.1. Plant growth

There were significant statistical interactions ðP  0:05Þ between time and temperature
for all plant growth variables, indicating that growth rate varied among the different
temperature treatments. Therefore, plant growth rates during the experiment are reported
separately at each temperature. Also, the relationship between root temperature and
plant height, stem diameter, shoot growth, the number of nodes per shoot, leaf chlorosis,
leaf abscission and leaf dry and fresh weights are shown only for the last measurement date
(day 42).
Trees grown at 25 or 38 8C had the fastest growth rate and greatest increase in plant
height, whereas trees grown at the 5 or 10 8C root temperatures had the slowest rate of

Fig. 2. Effect of root temperature on (a) plant height, and (b) stem diameter of 1-year-old, container-grown
‘Arkin’ carambola trees 42 days after root temperature treatments began. Each data point is the mean of six trees
from each of three replications per treatment. The regression line for (a) plant height is represented by
y ¼ 1:56 þ 6:41x, r2 ¼ 0:73, and (b) stem diameter by y ¼ 0:03 þ 0:36x, r2 ¼ 0:70. For both equations,
P  0:01.
72 H.L. George et al. / Scientia Horticulturae 96 (2002) 67–79

Fig. 3. Effect of root temperature on the increase in (a) axillary shoot length, and (b) the number of axillary
nodes per shoot of 1-year-old, container-grown ‘Arkin’ carambola trees 42 days after root temperature
treatments were initiated. Each data point is the mean of three shoots from each of six trees from three
replications. The regression line for (a) axillary shoot growth (length) is represented by y ¼ 0:03 þ 0:36x,
r 2 ¼ 0:49, and (b) the number of axillary nodes per shoot by y ¼ 0:19 þ 2:32x, r2 ¼ 0:42. For both equations,
P  0:01.

growth and the least increase among the other treatments (Fig. 1a). Plant height increased
significantly with an increase in root temperature (Fig. 1a). Stem diameter increased at a
greater rate for trees grown at the 38 8C root temperature than for trees at all other root
temperatures (Fig. 1b). The growth rate and final increase in stem diameter of trees growing
at 25 8C were relatively similar to those at 10 or 20 8C but greater than trees growing at the
5 8C root temperature. Moderate (20–25 8C) or low (5–10 8C) root temperatures had a
greater effect on plant height (Fig. 2a) than stem diameter (Fig. 2b).
As root temperature increased, axillary shoot length (Fig. 3a) and the number of nodes
per axillary shoot (Fig. 3b) increased significantly. The rate of axillary shoot growth
(Fig. 4a) and the rate of development of axillary nodes per shoot (Fig. 4b) was greatest for
trees exposed to a root temperature of 25 8C. Trees in the 38 8C treatment had a greater rate
of axillary shoot growth, and faster development of axillary nodes per shoot than trees at
the 5, 10, or 20 8C root temperatures.
The percentage of leaf chlorosis (Fig. 5a) and leaf abscission (Fig. 5b) increased
significantly as root temperatures decreased, although the coefficient of determination
between root temperature and these variables was not very high (r 2 ¼ 0:41 and 0.29 for leaf
chlorosis and abscission, respectively) (Fig. 5a and b). The rate of increase of leaf chlorosis
was greatest for trees exposed to root temperatures of 5 or 10 8C, intermediate for trees at
 H.L. George et al. / Scientia Horticulturae 96 (2002) 67–79 73

Fig. 4. Effect of root temperature on the increase in (a) axillary shoot length, and (b) the number of axillary
nodes per shoot, and internode length of 1-year-old, container-grown ‘Arkin’ carambola from 3 February to 8
June 1999. Each data point is the mean of three shoots from each of the six trees from each of the three
replications. Vertical bars indicate 1 S.E. Measurements were made at 14-day intervals. The regression lines
for (a) axillary shoot growth over time are represented by the equations: for 5 8C, y ¼ 0:42 þ 0:09x, r 2 ¼ 0:41;
for 10 8C, y ¼ 1:6 þ 0:22x, r2 ¼ 0:47; for 20 8C, y ¼ 1:18 þ 0:48x, r2 ¼ 0:48; for 25 8C,
y ¼ 2:76 þ 0:65x, r2 ¼ 0:47; for 38 8C, y ¼ 1:25 þ 0:44x, r 2 ¼ 0:68. The regression lines for (b) the
number of axillary nodes per shoot over time are represented by the equations: for 5 8C, y ¼ 0:12 þ 0:04x,
r2 ¼ 0:38; for 10 8C, y ¼ 1:14 þ 0:13x, r2 ¼ 0:49; for 20 8C, y ¼ 1:41 þ 0:18x, r 2 ¼ 0:50; for 25 8C,
y ¼ 3:58 þ 0:38x, r2 ¼ 0:53; for 38 8C, y ¼ 1:89 þ 0:26x, r 2 ¼ 0:56. For all equations, P  0:01.

the 38 8C root temperature, and least for trees growing at root temperatures of 20 or 25 8C
(Fig. 6a). The rate of leaf abscission was greatest for trees at a root temperature of 5 8C,
intermediate for trees at 10 or 38 8C and least for trees growing at 20 or 25 8C (Fig. 6b).
The response of leaf fresh and dry weight to root temperature was non-linear (Fig. 7).
Leaf fresh and dry weight increased with root temperature reaching a maximum at 25 8C
and decreased at 38 8C. The responses of stem and root fresh and dry weights to root
temperatures were not significant (data not shown). Fresh and dry matter accumulation was
generally greater in roots than in leaves or stems.

3.2. Leaf mineral element concentrations

There were no significant differences in initial leaf nutrient element concentrations

measured on day 0 among the root temperature treatments (data not shown). However, 42
days after root temperature treatments were imposed, leaf Ca, Mg, Fe, and Zn concentra-
74 H.L. George et al. / Scientia Horticulturae 96 (2002) 67–79

Fig. 5. Effect of root temperature on percent (a) leaf chlorosis, and (b) leaf abscission of 1-year-old, container-
grown ‘Arkin’ carambola trees 42 days after root temperature treatments were initiated. Each data point is the
mean of three trees from each of the three replications per treatment. The regression line for (a) leaf chlorosis is
represented by y ¼ 108:1013:90x, r 2 ¼ 0:41, and (b) leaf abscission by y ¼ 81:6711:27x, r 2 ¼ 0:29. For both
equations, P  0:01.

tions were significantly affected by treatment temperatures (Fig. 8). Total leaf nitrogen
concentration measured on day 42 of the experiment was not significantly different among
treatments and ranged from 28.0 to 37.8 g/kg dry wt. There were no significant differences
in leaf P and K among the five treatments on day 42 and their concentration ranged from 2.3
to 2.8, and 25.1 to 31.3 g/kg for P or K, respectively. There were no significant differences
in leaf Mn (20.8–65.2 g/kg), Cu (3.5–10.0 g/kg), and B (32.0–41.3 g/kg) concentrations
among treatments after 42 days (data not shown). In contrast, leaf Ca, Mg, Fe, and Zn
concentrations decreased sharply as root temperature increased from 5 to 10 8C, than
remained relatively constant from root temperatures of 10 to 38 8C (Fig. 8).

4. Discussion

Tree growth increased as root-zone temperature increased to 38 8C. Although carambola

trees can grow and produce fruit in subtropical environments, they are best adapted to
tropical environments, as indicated by the higher yields and better fruit quality in tropical
than in subtropical environments (Marler et al., 1994). In tropical environments, roots may
be exposed to temperatures between 25 and 38 8C. Thus, it is not surprising that carambola
growth increased as root temperatures increased up to 38 8C.
 H.L. George et al. / Scientia Horticulturae 96 (2002) 67–79 75

Fig. 6. Effects of root temperatures on leaf chlorosis and abscission of 1-year-old container-grown ‘Arkin’
carambola trees. Each data point is the mean of 10 leaves from each of the six trees from each of the three
replications per treatment. Vertical bars indicate 1 S.E. Measurements were made at 14-day intervals. The
regression lines for (a) leaf chlorosis over time are represented by the equations: for 5 8C, y ¼ 18:27 þ 2:29x,
r2 ¼ 0:75; for 10 8C, y ¼ 2:57 þ 2:33x, r 2 ¼ 0:87; for 20 8C, y ¼ 0:13 þ 0:97x, r2 ¼ 0:72; for 25 8C,
y ¼ 2:63 þ 0:74x, r 2 ¼ 0:71; for 38 8C, y ¼ 0:50 þ 1:52x, r 2 ¼ 0:81. The regression lines for (b) leaf
abscission over time are represented by the equations: for 5 8C, y ¼ 7:50 þ 2:05x, r2 ¼ 0:70; for 10 8C,
y ¼ 2:73 þ 1:51x, r 2 ¼ 0:79; for 20 8C, y ¼ 3:13 þ 0:62x, r2 ¼ 0:38; for 25 8C, y ¼ 2:80 þ 0:56x,
r2 ¼ 0:55; for 38 8C, y ¼ 1:97 þ 2:00x, r2 ¼ 0:77. For all equations, P  0:01.

In an orchard in southern Florida, mulching 5-year-old ‘Arkin’ carambola trees with

wood covered with clear polyethylene increased shoot growth, canopy density, and crop
yields compared to non-mulched trees (George et al., 2001c). Root-zone soil temperatures
for the mulched trees remained at or above 20 8C and averaged 25 8C, whereas root-zone
temperatures of the non-mulched trees were often below 20 8C. Thus, from the present root
temperature studies, it appears that the increased growth, and possibly the higher yields
were due to higher root-zone soil temperatures for mulched trees in the field experiment.
We observed wilting and dieback of apical buds on some shoots of trees grown at the 5
and 10 8C root temperatures 5–7 days after root temperature treatments were imposed. This
may explain the decline in axillary shoot length and number of nodes per shoot in these
treatments. Also, the poor growth response observed at root temperatures of 5 and 10 8C
may be associated with reduced root function at these temperatures which limited water
uptake. Average leaf xylem water potentials were more negative for carambola trees at root
temperatures 5, 10 or 38 8C compared to those at root temperatures of 20 or 25 8C (George
et al., 2001b). For citrus, low temperatures were shown to reduce root hydraulic
76 H.L. George et al. / Scientia Horticulturae 96 (2002) 67–79

Fig. 7. Effect of root temperature on leaf fresh and dry weights of 1-year-old, container-grown ‘Arkin’
carambola trees. Each data point is the mean of six trees from each of the three replications per treatment.
Measurements were made at the completion of each replication for fresh and dry weight, respectively. The
regression lines for fresh weight (*) are represented by y ¼ 22:79 þ 61:56x  7:98x2, r2 ¼ 0:62 and for dry
weight (*) by y ¼ 3:38 þ 23:741x  3:08x2, r2 ¼ 0:78. For both equations, P  0:01.

Fig. 8. Effect of root temperature on leaf Ca, Mg, Fe, and Zn content of 1-year-old, container-grown ‘Arkin’
carambola trees 42 days after root temperature treatments were initiated. Each data point represents the mean
of six trees from each of three replications per treatment. The non-linear regression lines are represented
by the equations: Ca ¼ 6:27 þ 660:83 etemp , r2 ¼ 0:57; Mg ¼ 3:51 þ 223:13 etemp , r2 ¼ 0:61; Fe ¼ 53:70 þ
25034:84 etemp , r2 ¼ 0:88; Zn ¼ 45:69 þ 5046:87 etemp , r 2 ¼ 0:64. For all equations, P  0:01.
 H.L. George et al. / Scientia Horticulturae 96 (2002) 67–79 77

conductivity (Wilcox et al., 1983). The significantly high percentages of leaf chlorosis and
abscission that occurred in the trees grown at the 5, 10, or 38 8C root temperatures may
have been due the roots’ inability to absorb adequate water to sustain the existing foliage.
Based on the present controlled-root temperature studies, it appears that increased root-
zone temperatures in a previous mulching experiment (George et al., 2001b) may have
been responsible for reduced leaf chlorosis and abscission.
An increase in water stress for plants in the low root temperature treatments may explain
the low foliage fresh and dry weights in trees at the 5, 10, and 38 8C root temperatures.
Similar growth responses have been observed in glasshouse studies of other tropical and
subtropical woody species. For example, the plant height and stem circumferences of
container-grown mango seedlings were greater at root temperatures of 27 and 21 8C than at
32 8C (Yusof et al., 1969). Diczbalis and Menzel (1998) reported that in rambutan
seedlings, maximum shoot length, leaf and node production, and dry matter accumulation
occurred at root temperatures ranging from 20 to 29 8C compared to 15 and 38 8C.
The rate of nutrient release by controlled release fertilizers may be affected by media
temperature and could affect plant growth and nutrient uptake by differences in media
nutrient content and/or an osmotic affect. However, we do not believe that an osmotic effect
would have occurred in the present study because (1) plants were fertilized only one time
during each replication of the experiment and each replication lasted only 42 days; (2)
containers were irrigated to container capacity three times per week, most likely preventing
a build-up of residual nutrients in the media; (3) research with containerized citrus has
demonstrated that it takes nearly an year to show a difference in tree response to Osmocote
because it is slow release (Ferguson et al., 1988). Furthermore, nutrient analysis from the
experiment suggested that no nutrient deficiencies occurred.
There is limited information on leaf element concentration for carambola and so
comparison of the nutrient concentrations observed in our study with critical ranges
for carambola was not possible. Galán Saúco et al. (1993) reported data from leaves of non-
bearing branches of carambola trees grown in Brazil. They reported nutrient element
concentrations of 14.0, 1.2, 9.8, and 6.4 g/kg for N, P and K, Ca, and Mg, respectively, and
microelement levels of 170, 73, 20, <5, and 51 mg/kg for Fe, Mn, Zn, Cu, and B,
respectively. The higher N, P, and K concentrations in our study may be due to differences
in nutrient availability among soil and artificial media, environment, tree age, and cultural
practices of field grown versus container grown trees. In contrast Mg, Ca, Mn, Zn, Cu, and
B concentrations reported by Galán Saúco et al. (1993) were within the range found in the
present study.
Studies conducted with other tropical and subtropical woody species showed an
interaction between their nutritional status and temperature. Yusof et al. (1969) found
that leaf N and P content of mango and avocado seedlings were highest at a soil
temperature of 32 8C and lowest at a soil temperature of 27 8C. Diczbalis and Menzel
(1998) observed that leaf N and P, and stem P concentrations were lower in rambutan
seedlings grown at root temperatures of 28 and 38 8C compared to the concentrations of
those seedlings grown at 15 and 20 8C in a glasshouse for 12 weeks. In a carambola orchard
in south Florida, increasing root-zone soil temperatures as a result of mulching generally
had no effect on leaf macro- or micro-element content (George et al., 2001a). In contrast,
in our controlled root temperature studies, leaf concentrations of Ca, Mg, Fe and Zn
78 H.L. George et al. / Scientia Horticulturae 96 (2002) 67–79

decreased as root-zone temperatures increased. Dilution effect may be involved in this

observation. Since the growth rate and leaf dry and fresh weights of trees at 5 8C was less
than that of the other treatments they may have accumulated higher concentrations of these
elements in the leaves than trees in the other treatments. The lower Ca, Mg, Fe, and Zn leaf
concentrations of trees grown at 10, 25, and 38 8C may have been a dilution effect. In the
orchard, root-zone soil temperature did not fall below 19.8 8C, even for non-mulched trees
(George et al., 2001a,b,c). Thus, the reduced growth as a result of lower soil temperatures
for non-mulched trees was probably not sufficient to cause a dilution effect for Ca, Mg, Fe
and Zn in an orchard.
Carambola tree growth is increased as temperature increases from 5 to 38 8C. This
corroborates the observations in an orchard where increasing root temperature due to
mulching increased carambola tree growth and yield (George et al., 2001a,b,c). The effect
of temperature on carambola tree growth does not appear to be related to plant nutrient
status, but may be related to increased water stress that occurs in roots held at low
temperatures (George et al., 2002).

Acknowledgements

Florida Agricultural Experiment Station Journal Series No. R-08177. We thank Drs.
Yuncong Li and Carl Campbell for helpful insights and discussion throughout this study
and T.E. Marler and R.C. Ploetz for critical review of this manuscript. This project was
partially supported by the Tropical Fruit Growers of South Florida, Carambola Research
Committee.

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