Leaf Gas Exchange of Pineapple as Influenced by Fruit

T.E. Marlera
Western Pacific Tropical Research Center
University of Guam
Mangilao, Guam 96923
USA

Keywords: Ananas comosus, crassulacean acid metabolism

Abstract
‘Smooth Cayenne’ pineapple plants were propagated from suckers and
grown in containers for leaf gas exchange studies. Following natural flowering and
fruit development, net CO2 exchange was measured in the leaves within the crown,
slips, and the main plant throughout diel cycles. Maximum net CO2 exchange of
leaves on the main plant was 0.44 μmol·m-2·s-1 and did not differ among vegetative
plants, flowering plants, and plants supporting fruits. Maximum net CO2 exchange
of crown leaves was 4.39 μmol·m-2·s-1 and that of slip leaves was 3.99 μmol·m-2·s-1.
The results indicate leaves subtending pineapple fruits do not increase in carbon
assimilation during fruit growth as do leaves of other fruit species. Based on these
gas exchange data, crown leaves may satisfy most of the carbon requirements of the
developing pineapple fruit on intact plants.

INTRODUCTION
Inflorescence induction in pineapple generates an apical reproductive structure. In
the typical plant with determinate flowering, the developing fruits are supported
exclusively by the pre-existing subtending leaves. However, in pineapple the
inflorescence and developing fruit are always augmented with new photosynthetic leaves
in developing crowns and sometimes augmented with leaves of slips (Bartholomew and
Malézieux, 1994). The sink activity due to presence of fruit generally increases net
photosynthesis of leaves near the fruit (e.g., DeJong, 1986; Lenz, 1979; Mondal et al.,
1978; Schaffer et al., 1986, 1987). The present study examined how flowering and the
presence of developing fruit on pineapple plants influenced net CO2 exchange and diel
gas exchange patterns in various types of leaves.

MATERIALS AND METHODS

Experimental Units
Experimental units were created by planting ‘Smooth Cayenne’ pineapple suckers of
uniform size in 5.1-L plastic containers. Plants were grown in peat:perlite (70:30%) medium.
Each plant received 100 ml of half-Hoagland’s solution weekly and daily watering when
needed.
Natural flowering of many plants occurred after 13 months of growth. This provided
an opportunity to study accumulation and partitioning of dry matter in fruiting and non-
fruiting plants of uniform age. Fruiting and vegetative plants were arranged in a Randomized
Complete Block design during fruit development. The plants were grown until color break
on the fruits. This provided an opportunity to study leaf physiology as influenced by
presence or absence of flowers and developing fruits.

Gas Exchange Measurements

Leaf gas exchange measurements were recorded using a CIRAS-1 portable
photosynthesis system (PP Systems, Stotfield, Hitchen, Herts., UK). For each measurement,
a fully expanded leaf was inserted into the system’s 2.5-cm2 leaf cuvette. In order to define
a
tmarler@uguam.uog.edu

Proc. 7th International Pineapple Symposium 239

Eds.: H. Abdullah et al.
Acta Hort. 902, ISHS 2011
the best time of day to compare gas exchange among the leaf types, net CO2 exchange
was measured approximately every two hours throughout entire diel cycles. This was
done on several days at flowering, during the rapid growth phase of the fruit, and at the
end of the study when fruits began showing color. Net CO2 exchange was measured on
leaves of the main plant beginning at the flowering stage and continuing through fruit
harvest. Net CO2 exchange was measured on crown leaves during the second half of
crown development. Some of the reproductive replications contained slips at the apex of
the peduncle. Therefore, I also compared leaf gas exchange on plants with and without
slips.

RESULTS AND DISCUSSION

The shape of the diel pattern of net CO2 exchange did not differ between fruiting
and vegetative plants during the flowering stage (not shown). The general shape of the
diel pattern of gas exchange of main plant and crown leaves did not differ during the main
fruit growth stage, when temperature ranged from 26-35°C and ambient CO2 ranged from
351-355 μl/L (Fig. 1). The daily pattern of gas exchange also did not differ among crown
and main plant leaves when fruits were mature (not shown). For plants with slips, the
daily pattern of gas exchange did not differ among leaves in the crown, slips, or the main
plant. Net CO2 efflux during the diurnal phase was not different among all measurement
days and among all leaf types. Shortly after sunset, differences in net CO2 exchange
among leaf types emerged and reached a maximum in all cases during Phase I of the
CAM cycle from 2200 h through 0200 h.
Maximum net CO2 exchange of leaves of flowering plants did not differ from that
of leaves of vegetative plants throughout the entire flowering stage (Fig. 2). Maximum net
CO2 exchange of crown leaves during Phase I of the CAM cycle greatly exceeded that of
leaves on the main plant (P=0.0003, Fig. 3). However, maximum net CO2 exchange of
crown leaves did not differ between reproductive plants with or without slips (not shown).
In reproductive plants with slips, maximum net CO2 exchange of leaves differed
among main plants, slips, and crowns (P=0.0014, Fig. 3). Net CO2 exchange of leaves
from slips was not different from that of leaves from crowns. Temperature from 2200-
0200 h during these comparisons of different leaf types was 25-26°C and ambient CO2
ranged from 350-356 μl/L.
The source-sink relations of developing fruit and leaves near those fruits have
been well-studied for many angiosperms. As a general rule, leaves near fruits increase in
photosynthetic activity during the demanding stages of fruit development. In this study,
pre-existing pineapple leaves did not adjust their photosynthesis in response to increased
sink activity of flowering or fruiting. This lack of response may be due to the production
of new photosynthetic tissue in the form of the crown and sometimes the slips. This new
leaf surface may satisfy the increased carbon needs of the plant that result from
inflorescence induction and fruit development.
Pineapple plant morphology does not allow the use of some traditional methods
for manipulating source-sink relations. For example, fruit removal (e.g., DeJong, 1986;
Mondal et al., 1978; Schaffer et al., 1986, 1987) would not be informative because crown
leaves cannot be left intact during the process of fruit removal. The combination of
girdling and selective defoliation of stem sections to block import of substrates from
source leaves to fruits (e.g., Hoch, 2005) is also impractical in pineapple because of the
location of phloem within peduncle and the lower central axis of the crown.
Several reports indicate that complete or partial crown removal does not influence
fruit size or increases pineapple fruit size (Chen and Paull, 2001; Hwang, 1968; Prakash
et al., 1983; Wee and Ng, 1970). These results do not corroborate the conclusion that
crown leaves satisfy the needs of developing fruits. However, compensatory source
activity in leaves of the main plant that results from crown removal may account for the
lack of influence on fruit size, and these results may have little application to
understanding carbon relations in an intact plant.
Light load during the photoperiod influences pineapple leaf net CO2 exchange

240
during subsequent nocturnal portions of the CAM cycle (Bartholomew and Malézieux,
1994; Nose et al., 1977; Sale and Neales, 1980). Moreover, shading pineapple plants
reduces total soluble solids in fruits (Py et al., 1987). Therefore, manipulating the light
load of crown, slips, and/or basal leaves in various combinations and measuring leaf gas
exchange during the following nocturnal period may clarify some of the source-sink
questions that remain unanswered. In some production areas, shading is used to reduce
sunburn on fruit tissue (Broadley et al., 1993). Results from this study indicate this
practice may reduce crown leaf performance.
Temperature variations influence several relevant factors such as leaf area ratio
and development of the crown, suckers, and slips in relation to fruit (Bartholomew and
Malézieux, 1994). Therefore, adjusting temperatures during fruit development to
manipulate source-sink relations may be another approach to improving our
understanding of carbon relations in this unique plant. Ultimately, incubating leaves of
crown, slip, or the main plant with 14CO2 and then quantifying the percentage of labeled
photosynthate among the organs is the only unambiguous means of clarifying the source-
sink relations of an intact, fruiting pineapple plant.

Literature Cited
Bartholomew, D.P. and Malézieux, E.P. 1994. Pineapple. p.243-291. In: B. Schaffer and
P.C. Andersen (eds.), Handbook of Environmental Physiology of Fruit Crops, Vol. II.
Sub-tropical and Tropical Crops. CRC Press, Boca Raton, Florida.
Broadley, R.H., Wassman, III, R.C. and Sinclair, E.R. 1993. Pineapple Pests and
Disorders. Queensland Dept. Primary Ind., Brisbane.
Chen, C.C. and Paull, R.E. 2001. Fruit temperature and crown removal on the occurrence
of pineapple fruit translucency. Scientia Hortic. 88:85-95.
DeJong, T.M. 1986. Fruit effects on photosynthesis in Prunus persica. Physiol. Plant.
66:149-153.
Hoch, G. 2005. Fruit-bearing branchlets are carbon autonomous in mature broad-leaved
temperate forest trees. Plant, Cell & Environment 28:651-659.
Hwang, Y.J. 1968. The effect of removing the crowns on the quality and yield of
pineapples. J. Agric. Assoc. China 62:48-62 (Chinese, with English summary).
Lenz, F. 1979. Fruit effects on photosynthesis, light- and dark-respiration. p.271-281. In:
R. Marcelle, H. Olljsters and M. Van Poucke (eds.), Photosynthesis and Plant
Development. Dr. W. Junk Publishers, The Hague, London.
Mondal, M.H., Brun, W.A. and Brenner, M.L. 1978. Effects of sink removal on
photosynthesis and senescence in leaves of soybean (Glycine max L.) plants. Plant
Physiol. 61:394-397.
Nose, A., Shiroma, M., Miyazato, K. and Murayama, S. 1977. Studies on dry matter
production in pineapple plants. I. Effects of light intensity in light period on the CO2
exchange and CO2 balance of pineapple plants. Japan. J. Crop Sci. 46:580-587.
Prakash, G.S., Reddy, B.M.C. and Dass, H.C. 1983. The effect of partial pinching of
crown at different stages of its growth on the size and shape of fruit in ‘Kew’
pineapple. Singapore J. Primary Indust. 11:101-105.
Py, C., Lacoeuilhe, J.J. and Teisson, C. 1987. The Pineapple, Cultivation and Uses.
Maisonneuve et Larose, Paris.
Sale, P.J.M. and Neales, T.F. 1980. Carbon dioxide assimilation by pineapple plants,
Ananas comosus (L.) Merr. I. Effects of daily irradiance. Aust. J. Plant Physiol. 7:363-
373.
Schaffer, B.J., Barden, J.A. and Williams, J.M. 1986. Net photosynthesis, stomatal
conductance, specific leaf weight, and chlorophyll content of strawberry plants as
influenced by fruiting. J. Amer. Soc. Hort. Sci. 111:82-86.
Schaffer, B., Ramos, L. and Lara, S.P. 1987. Effect of fruit removal on net gas exchange
of avocado leaves. HortScience 22:925-927.
Wee, Y.C. and Ng, J.C. 1970. Decrowning and deslipping of the Singapore Spanish
pineapple. Trop. Agric. (Trinidad) 47:73-75.

241
Figures

Fig. 1. The influence of time of day on net CO2 assimilation for ‘Smooth Cayenne’
pineapple leaves in the crown and main plant.

Fig. 2. Maximum net CO2 exchange of leaves on the main plant during Phase 1 of the
CAM cycle for vegetative plants, reproductive plants during the flowering stage,
and fruiting plants as fruits mature. Bars are standard error.

242
Fig. 3. Maximum net CO2 assimilation of leaves on the main plant (base), the crown, and
the slips of fruiting ‘Smooth Cayenne’ pineapple plants during Phase 1 of the
CAM cycle. Bars are standard error.

243
244