Ann. appl. B i d .

(1993), 123,221-232
Printed in Great Britain 221

Flower and nectar attributes of pepper (Capsicum

annuum L.) plants in relation to their attractiveness to
honeybees (Apis mellifera L.)
By H D RABINOWITCH', A FAHN*, TAL MEIR and Y LENSKY
The Hebrew University of Jerusalem, Faculty of Agriculture, Rehovot
76 100, Israel
* The Hebrew University of Jerusalem, Institute of Life Sciences, Department
of Botany, Giva't Ram, Jerusalem 91904, Israel
(Accepted 29 January 1993)

Summary
Flower morphology, nectary structure, nectar traits and rates of honeybee
foraging on pepper plants were studied. The nectary appears as swellings on the
basal part of the ovary. The nectariferous cells are smaller and denser than the
neighbouring parenchyma. Stomata are present in the nectary epidermis, but do
not appear on the other parts of the ovary epidermis. Seven pepper breeding
lines were grown near a bee yard in Rehovot. Five to six fold differences in
nectar volume were found between the extreme genotypes. Nectar volumes were
higher during noon and afternoon hours, as compared with morning hours. High
correlation coefficients between nectar volume and sugar concentration were
found. These were significant for the four high nectar yielding genotypes, ranging
between r = 0.65 to r = 0.94. Male-fertile flowers produced more nectar and
higher sugar concentration than sterile ones. Skewed distribution was observed
in nectar volume of F2 populations, but relatively low heritability values were
calculated.
Pepper nectar contains fructose and glucose only. The former occupies 52-
82% of the total sugar content. Pepper genotypes varied in frequency of honeybee
visits and significant correlation between sugar quantity and number of honeybee
visits per flower was evident. Fertile pepper flowers are not very attractive to
honeybees and male-sterile flowers are even less so. The considerable variation
in nectar characteristics can be exploited to increase attractiveness to honeybees,
thus facilitating bee pollination in commercial production of F1hybrid seeds and
improve fruit quality.

Key words: Pepper, Capsicum annuum, honeybees, Apis mellifera, nectaries,

nectar characteristics, seed production, male-sterility, floral charac-
teristics

Introduction
There is little information on the pollination of sweet pepper (Capsicum annuum L.) and
this tends to be contradictory (Free, 1970, 1975; McGregor, 1976; Pouvreau, 1984).
McGregor (1976) stated that peppers and other members of the Solanacae are noted for
their low attractiveness to honeybees. On the other hand, many researchers suggested that
honeybees (Rylski, 1986), thrips and ants play a role in cross-pollination of these flowers
'Corresponding author
0 1993 Association of Applied Biologists
222 H D RABINOWITCH, A FAHN, TAL WEIR AND Y LENSKY

(Erwin, 1932; Hawthorn & Pollard. 1954; Markus, 1965; Odland & Porter, 1941), but only
10-30% cross-pollination was reported (Murthy & Murthy, 1962; Odland & Porter, 1941;
Purseglove, 1968).
Honeybees are attracted to pollen and nectar (von Frisch, 1967), and especially to nectar
sugars (Percival, 1961). The only report known to us on sugar composition of pepper nectar
indicated the presence of glucose only (Martin, Erwin & Lounsberry , 1932). but the presence
of sucrose was not tested. No information is available on daily nectar fluctuations in pepper
flowers and on the genetic variation in nectar volume and in sugar contents. A number of
workers reported male sterility in peppers (Martin & Crawford, 1951; Peterson, 1958;
Shifriss & Frankel, 1969). This trait is currently being exploited by seed companies for F,
seed production. No comparison was made, however, on nectar attributes between male-
sterile and -fertile pepper flowers.
According to Martin et al. (1932) the nectar in the genus Capsicum is secreted from the
base of the corolla tube and the adnated portions of the stamen filaments. However, in
most Solanaceae species the nectariferous tissue has been reported to be associated with
the ovary (Bernardello, 1987; Bonnier, 1879; Daumann, 1928).
We hereby present our observations on the position, structure and morphology of pepper
flower nectaries, as well as on the variation in nectar characteristics of pepper. The relation
between nectar traits and frequency of honeybee visitation was also investigated and the
implication on hybrid seed production is discussed.

Materials and Methods

Plant material
Seven sweet pepper ( C . annuum L . ) breeding lines differing in fruit shape, size and taste
were used, as detailed in the Results; so were fertile and male-sterile isolines of a further
two breeding lines.

Experimental
During the summer months of 1986 to 1988, individual pepper seedlings were transplanted
at random into 25 m long rows at a spacing of 50 and lOOcm between plants and rows,
respectively; there were about 25 plants per parental line and F,, and 100 to 200 plants per
segregating population in experimental plots, as detailed in the Results. These were located
50 m from a beeyard of 80 Italian honeybee (Apis rnellifera L. var. figustica) colonies at the
Triwaks Bee Research Center, Rehovot.

Preparation of sections f o r microscopy

Freshly harvested flowers were fixed in formalin-acetic acid (FAA) and dehydrated in a
graded series of ethanol. Some of the dehydrated material was embedded in paraffin
(Jensen, 1962), and slices 8 p n thick stained with alcian green-safranin (Joel, 1983). Other
dehydrated flowers were critical point dried with solvent-substituted liquid carbon dioxide,
coated with a thin layer of gold and examined with a Joel JC M35 scanning electron
microscope.

Bee visitation
The number of honeybees visiting pepper flowers per plant was counted while steady
strolling along the row for 15 min and all bee visits to open flowers were recorded. These
 Pepper power attributes and attractiveness to honeybees 223
counts were made at 15 min intervals (A). The number of open flowers per plant (B) was
counted every morning. Number of visits per flower per plant was calculated as A/B. For
each genotype, this procedure was repeated three times during the experimental season
between 8.30 am and 2.00 pm.

Nectar
Nectar was collected between 8.00 am and 2.30 pm. One day before anthesis, flower buds
were covered with gelatin capsules (0.8 cm X 2.0 cm). Flowers were picked at anthesis, and
nectar was collected immediately thereafter. Five or 10 p1 graduated glass capillaries (Brand,
Wertheim, Germany) and a portable refractometer (Optinat, model 6351, range 0-32%,
Tokyo, Japan), were used to determine the nectar volume and sugar concentration,
respectively. Pooled nectar samples were stored in microfuge tubes at - 18"C, until used.

Heritability
The heritability of nectar volume and sugar concentration was calculated according to
Falcooner (1981) as follows:

hZb= [Var(F2)- 0.25Var(P,)+ 0.25Var(P2)+OSVar(F,)]/Var(FJ

PI and P2 were the respective female and male parental lines of the F1 and F2 offspring
populations.

Sugar composition
Nectar analysis for fructose, glucose and sucrose were performed as follows: After 24 h
lyophilisation, nectar samples were treated with 20 pl Sil-A (trimethyl sylil, Sigma, St Louis,
MO, USA) per 1 mg dry nectar for 60 min at room temperature. One pl aliquots were
injected into Packard model 7400 gas chromatograph using glass columns (2 m long, i.d.
4 mm) packed with 3% SE-30 (Supelco, Bellefonte, PA, USA). The separation conditions
were as follows: injection port and detector 250°C; oven temperature was programmed
from 110-285°C at 3"C/min or 5"C/min. The flow of gases (cc/min) was as follows: N,-40
or 70; H2-40; Air-400. &-fructose, a-glucose, b-glucose and sucrose (Sigma) were used as
standard sugars.
Coefficient of variation was less than 15% in all experiments.

Results

Floral morphology
The corolla of the pepper flower is gamopetalous, rotate, or nearly so, and lobed. The
basal portion of the corolla is in the form of a short tube. The stamens alternate with the
petal lobes. The bases of the filaments of the stamens are adnated to the corolla tube. The
separation of the filaments from the corolla tube takes place a short distance below its upper
rim (Figs 1-4). At this position projections occur on both sides of the widened basal part
of each filament (Figs 2 and 5). There is a deep groove extending almost to the very base
of the corolla tube between the filament bases and their projections (Figs 2, 5 and 6). The
ovary exhibits at its base a swelling, which varies in height at various places around the
ovary (Figs 7 and 8). This swelling represents the nectary. Its cells are smaller and denser
than other parenchymatic cells of the ovary wall and of the parenchyma cells of the corolla-
224 H D RABINOWITCH. A FAHN, TAL MEIR AND Y LENSKY

Ca
CT
SF
0

Figs. 1-4. Diagrams of a pepper flower sectioned at various positions. Fig. 1. Longitudinally split flower.
Fig. 2. Portion of a corolla with the adnated stamens. Fig. 3. Cross-section at the top of the corolla
tube. Fig. 3 . Cross-section of the corolla tube; the filament bases are adnated to the corolla tube.
Ca - calyx; C F r - corolla tube with the adnated filament bases; CL - corolla lobes; SF - stamen
filaments; FB - filament bases; 0 - ovary; N - nectary; Pr - projections of the widened filament bases;
R - flower receptacle; S - stigma.

filament tube (Fig. 7). Application of Fehling’s solution (Bonnier, 1879), results in colour
change of the region of these cells to orange-brown, thus indicating the presence of reducing
sugars. The cells below the nectariferous tissue are many fold larger than the nectarial cells
and the rest of the ovarial wall (Fig. 8). There is a clear distinction between the border of
the ovary wall, including the nectariferous tissue, and the flower receptacle. The cells of
the latter are relatively very large and elongated in the radial direction. Stomata are present
only in the epidermis of the nectary region (Fig. 9), but not in the rest of the ovary. The
nectary stomata are widely open (Fig. lo), and irregularly distributed (Fig. 9).
 Pepper flower attributes and attractiveness to honeybees 225

Figs. 5-8. Electron micrographs of floral organs. Fig. 5 . Scanning electron micrograph (SEM) of stamens
with projections (Pr) on the edges of the widened filament bases. F - stamen filament. Scale bar =
500 pm (X40). Fig. 6. Portion of a flower cross-section at the level where the stamen filaments are
adnated to the corolla tube. Grooves (G) are present between the adnated basal parts of the filaments
-
(F). Ca - calyx; CT - corolla tube; N nectariferous tissue; 0 - ovary. Scale bar = 250 km ( ~ 8 7 )Fig.
.
7. SEM of an ovary. N - nectary. Scale bar = 400 pm ( ~ 5 0 )Fig. . 8. Partial longitudinal section of a
flower showing the nectariferous tissue (N) at the base of the ovary wall (OW). Ca - calyx; CT - corolla
tube with adnated filament base; F - stamen filament; R - flower receptacle. Scale bar = 250 pm (~85).

Diurnal changes in nectar secretion and genorypic differences

Nectar volume per flower of seven pepper genotypes was markedly higher around noon
and early afternoon than in the morning. Five to six fold differences in nectar volume were
found between the extreme types on a daily basis. Ranking order varied between sampling
times, but the extreme genotypes maintained their positions (Fig. 11).Nectar volume and
226 H D RABINOWITCH. A FAHN. TAL MEIR AND Y LENSKY

Figs. 9-10, SEM micrographs of ovarial and nectary tissues. Fig. 9. Ovarial portion: Stomata (St) occur
only at the base of the ovary. i.e. in the region of the nectary ( N ) . Scale bar = 100 pm ( ~ 2 2 0 )Fig.
. 10.
Modified stoma of the nectary with a widely open aperture. Scale bar = 10 pm ( x2400).

\ugar quantity were closely correlated in the four high nectar yielding genotypes. with
c.oefficients of 0.73.0.65,0.87 and 0.94 for H, A Y , AS and AP, respectively, but not in the
poor yielding SY. KS and KH genotypes. Correlation between nectar volume and sugar
concentration was low (data not shown). These results indicate the presence of genetic
variation for nectar secretion in C. anmiurn and the possibility of selecting for large nectar
isulume with no effect on sugar quantity.
 Pepper flower attributes and attractiveness to honeybees 227

Genotypes

9 :00-10 :30 11 :W 1 2 :30 13 : Mk I4 : 30

Sampling time, h
Fig. 11. Daily pattern in nectar volume production of seven pepper genotypes.

Comparisons between nectar traits of male-sterile and male-fertile isolines

Nectar collections were made from male-fertile (mf) and male-sterile (ms) isolines ( F6
populations). Between 9.00 and 10.30 am, mf flowers produced significantly higher volume
of nectar and larger amounts of sugars than ms flowers, but at no other time of day (Table
1). Sugar concentration in the nectar increased throughout the day in both the ms and mf
flowers, but with no significant differences between the two genotypes.

Heritability
Nectar was collected from two parental lines and from their F1and F2offspring populations
(Table 2). Skewed distribution of the F2 population shown in Fig. 12 indicates the presence
of genotypes excelling in nectar secretion. However, the low heritability value for nectar
volume of h2, = 0.12, indicates that selection for high nectar volume requires screening of

Table 1. Pepper nectar volume. sugar concentration and quantity in two sets of male fertile
and sterile pepper isolines. Coefficient of variation was less than 15% in all cases
Male-Fertile Male-Sterile
flowers flowers
Nectar Sugar Sugar Nectar Sugar Sugar
volume conc. quantity volume conc. quantity
Hour (Pi) (%) (10-2 mg) (Pl) (%) (10-2mg)
Genotype BR
10 8.6 4.9 0.42 6.8 5.3 0.36
12 8.8 6.8 0.60 7.4 7.0 0.52
I4 6.9 7.3 0.50 6.8 7.3 0.50
Genotype MA
10 1.7 23.9 0.41 1.0 22.2 0.22
12 1.3 24.6 0.32 I .3 21.2 0.28
228 H D RABINOWITCH, A FAHN. TAL MEIR AND Y LENSKY

24

>? 18
c
1
W
2
LL 12

0-1 1-2 2-3 3-4 4-5 5-6 6-7 7-8 8-9 9-10 10-11 11-1212-1313-14 14-15
Nectar volume. ptl/flower

Fig. 12. Nectar volume distribution of F? (Maor X Spanish) population. Nectar was collected in the
morning hours. Population size = 208 plants.

large populations under a variety of environmental conditions. No differences in sugar

concentration were found between the parental lines, therefore no estimates of heritability
could be made for this trait.

Sugar composition
Nectar collected from four non-related pepper genotypes was analysed for sugar com-
position using gas chromatography. Fructose and glucose were the only sugars detected,
and fructose was the major constituent (Fig. 13).

Frequency of honeybee visits

Frequency of honeybee visits per flower was relatively high in SY and AP genotypes, as
compared with that in KS and AS (Fig. 14). There was a significant correlation between
sugar quantity and number of bees' visits per flower, both in the morning and afternoon
hours (r = 0.62 and 0.73, respectively), but not for nectar volume and sugar composition.
The foraging activities on fertile and sterile flowers were: 18 5 ' 3 and 4 -e 2'visits per 100
flowers per 15 min, respectively.

Table 2. Nectar volume and sugar concentration in pepper flowers of parental lines and the
F, and F, populations
Nectar volume, pl Sugar concentration, 76
Flower Flower
Population number Mean number Mean
p, 111 3.3b' 127 7.h
pz 31 7.2a 29 7.1a
F, 24 5.7ah 21 6.4,
F2 1191 5.2ab 968 h.3a

*Means followed by the same letter do not significantly differ at P = 0.05.

 Pepper flower attributes and attractiveness to honeybees 229

= Fructose @@ Glucose

AP AS SY KS
Genotype

Fig. 13. Fructose and glucose levels in nectar of four pepper genotypes.

Discussion
In most Solanacea species, nectariferous tissue is located either at the base of the ovary,
usually as a swelling, or just beneath it, as an annular disc (Bernardello, 1987; Bonnier,
1879; Daumann, 1928; Galetto, 1991; Kartashova, 1965; Knuth, 1899). In Sofunum dul-
camara nectar is secreted from the basal projections of the stamen filaments (Knuth, 1899).
whereas in Capsicum spp. nectar is secreted from both the corolla tube and the adnated

0.40

0.30

0.20

0.10

I I I I I I I I I I I I

8 9 9:30 10 10:30 11 11:30 12 12:30 13 13:30 14 14:30

Hour

Fig. 14. Daily patterns of honeybee visitation per flower of four pepper genotypes. Observations started
at 8 AM, 10 AM and 1 PM and lasted 15 min, each.
230 H D RABINOWITCH, A FAHN. TAL MEIR AND Y LENSKY

portions of the filaments (Martin et al., 1932). Our data show that in C. annuurn nectar is
secreted solely from the swollen base of the ovary (Figs 7 and 8); the deep grooves in the
corolla-filament tube, conduct the nectar from the nectary towards the upper part of the
corolla tube, but do not secrete by themselves.
This variation in nectary location raises the question of the evolutionary changes in its
position. Fahn (1953) suggested that the migration of the nectary was mainly in an acro-
centripetal direction, from the perianth towards the ovary, style and in some cases to the
stigma. A study by Dauman (1928) showed that in mature flowers of Nicotiana glauca, the
nectary is situated on the ovary. and appears as a ring between the ovary and the flower
receptacle. This and the fact that in C. annuurn stomata occur only in the epidermis above
the nectariferous tissue but not in the rest of the ovarial epidermis, strengthens the view
that in Solanacae, the nectary originated in the region of the flower receptacle and shifted
during evolution to the ovary.
It is quite obvious that nectaries of pepper flowers are within easy access to honeybees
and other insects. Hence, nectar quality and/or other flower attributes affect the pepper
flowers' attractiveness to pollinators. Nectar volume is not the major factor restricting bee
visitation to pepper flowers. A high frequency of visits of honey bees was measured in
Labiatae flowers which secrete only about 0.31 pl of nectar per flower per day (e.g.
Rosmarinurn oficinalis and Coridothymus capitatus; Dafni, Lensky & Fahn, 1988). Indeed,
the frequency of bee visitation to pepper flowers did not change much with the increased
nectar secretion at noon-time. It is therefore concluded that the low attractiveness of
pepper nectar to honeybees is due to other floral factors, such as sugar composition and
concentration.
Honeybees do not usually collect nectar with less than 13% sugars unless water is needed
to dilute stored honey for brood rearing (Crane, 1990) or for cooling the broodnest during
summer (Lensky, 1964). Our results show that pepper nectar is often poor in sugars, with
concentrations ranging from 5.3-24.6%. This may explain the low attractiveness of pepper
flowers to honeybees and other insects and the consequent low rates of cross-pollination
(McGregor. 1976: Murthy & Murthy. 1962; Odland & Porter, 1941; Purseglove, 1968).
Martin et al. (1932) identified glucose only in C. annuurn nectar. Our results show that
fructose is dominant (Fig. 13). This discrepancy may be explained by the different genotypes
employed by the two groups. There is no agreement among researchers on the preference
of honevbees for various nectar sugars (Bachman & Waller, 1977: Estes, Amos & Sullivan,
1983; Jamieson & Austin, 1956; Percival. 1961: Waller. 1972; Wykes, 1952). However,
some workers reported that total sugar concentration is the main attracting factor to
honeybees (Waller. 1972).
Our results show that genotypes vary in nectar volume per flower and positive correlation
was found between nectar volume and sugar quantity. The latter, was highly correlated
with the frequency of bee visitation. Regardless of the low heritability. these results, taken
together with the skewed distribution of the F, population, indicate the possibility for
selecting improved plants more attractive to honeybees than the currently available geno-
tvpes.
It is well established that seed set is obligatory for pepper fruit development and that
partial seed set results in small malformed fruit (McGregor, 1976; Pouvreau, 1984). It was
estimated that 30% or less cross-pollination occurs in peppers. Therefore, any means for
increasing honeybee visitation could increase quality yields.
Male-sterile plants produce significantly lower volumes of nectar and smaller amounts of
sugars than fertile ones, at least early in the day. We have also noticed that bee visitation
to male-sterile flowers was 4-5 fold lower than that to the fertile flowers. Many modern
pepper cultivars are F, hybrids. Male-sterile plants are utilised as female parents, but
 Pepper f l o w e r attributes and attractiveness to honeybees 231

pollination is done manually. This is a labour-intensive task, thus increasing the cost of
hybrid seed production. Male-sterile flowers lack pollen, and therefore increased attractivity
to honeybees can be gained only by improving nectar characteristics. If this is achieved then
manual labour connected with hybrid seed production would be greatly reduced.
Our work shows large variation among pepper genotypes in nectar characteristics and in
attractivity to honeybees and therefore in the potential for improved seed set and fruit
quality through selection.

Acknowledgements
We thank Mr Serge Ben-Harush and Dr Yehiel Tal (Mivkhor Farm), Mr Jim Winkler
and Mr Zakhi Rotem (Berrurim Farm), Hazera Seed Company, for their helpful assistance.
We thank Ms Hani Gal for her excellent assistance in the preparation of the manuscript
and the figures. T Meir has been awarded a research fellowship by the Hazera Seed
Company and Benjamin and Gavriel Triwaks Fund.

References
Bachman W W, Waller G D. 1977. Honeybee responses to sugar solutions of different compositions.
Journal of Apicultural Research 16:16.5-169.
Bernardello L M. 1987. Comparative floral morphology in Lycieae (Solanaceae). Brittonia 39:112-
129.
Bonnier G. 1879. Les nectaires. Theses, Paris: Faculte des Sciences, Universite de Paris.
Crane E. 1990. Bees and Beekeeping. Oxford: Heinemann Newnes. 614 pp.
Dafni H, Lensky Y, Fahn A. 1988. Flower and nectar characteristics of nine species of Labiatae and
their influence on honeybee visits. Journal of Apicultural Research 27:103-114.
Daumann E. 1928. Zur Biologie der Blute von Nicotiana glauca Grah. Biologia Generalis 4571-588.
Erwin A T. 1932. The peppers. Bulletin of the Iowa Agricultural Experimental Station 293:12@152.
Estes J R, Amos B B, Sullivan J R. 1983. Pollination from two perspectives: the Agricultural and
Biological Sciences. In Handbook of Experimental Pollination Biology, pp. 536-554. Eds C E Jones
and R J Little. New York: Scientific & Academic Editions. Van Nostrand Reinhold Co. Inc.
Fahn A. 1953. The topography of the nectary in the flower and its phylogenetical trend. Phy-
tomorphology 3:424-426.
Falcooner D S, 1981. Introduction to Quantitiue Genetics. London: Longman.
Free J B. 1970. Insect Pollination of Crop Plants. London & New York: Academic Press. 544 pp.
Free J B. 1975. Pollination of Capsicum frutescens L., Capsicum annuum L. and Solanum melongela
L. (Solanacae) in Jamaica. Tropical Agriculture (Trinidad) 52:353-357.
Frisch K von. 1967. The Dance Language and Orientation of Bees. Cambridge, MA: The Belknap
Press, Harvard University Press. 565 pp.
Galetto L. 1991. Sobre el nectar y 10s nectarios de algunas especies de Nicotiana (Solanaceae).
Kurtziana 21:165-176.
Hawthorn L R, Pollard L H. 1954. Vegetable Seed and Flower Production. New York: The Blackiston
Co., Inc. 626 pp.
Jamieson C A, Austin G H. 1956. Preference of honeybees for sugar solutions. Proceedings of the
Tenth International Congress of Entomology, Montreal 4: 105S1062.
Jensen W A. 1962. Botanical Histochemistry. San Francisco: Freeman. 408 pp.
Joel D M. 1983. AGS (Alcian Green Safranin) - A simple differential staining of plant material for
the light microscope. Proceedings of the Royal Microscopic Society 18:149-151.
Kartashova N N. 1965. Stroeniei Funktsia Nektarnikou Tsvetua Duudol’nikh Rastenii. Tomsk:
Izdatel’stvo Tomskogo Universiteta. 194 pp.
Knuth P. 1899. Handbuch der Blutenbiologie. Volume 2, p. 128. Leipzig: Wilhelm Engelmann.
'32 H D RABINOWITCH, A FAHN. T A L MElR AND Y LENSKY

Lensky Y. 1964. Comportement d'une colonie d'abeilles a des temperatures extremes. Journal of
Insect Physiology 10:279-282.
Markus P. 1965. Investigation on cross-fertilization in red pepper for spice. Hungarian Agricultural
Reoiew 13:750 (In Hungarian).
Martin J A, Crawford J H. 1951. Several types of sterility in Capsicurn frutescence. Proceedings o f
the American Society of Horricirltural Science 57:335-338.
Martin J A, Erwin A T, Lounsberry C C. 1932. Nectaries of Capsicutn. lou,a State College, Joiirnal
of Science 6:277-285.
McGregor S E. 1976. Eggplant, Pepper. Tomato. In Insect Pollination of C'ultic~atedPlants. pp. 213-
215; 292-295; 357-361. Washington DC: USDA.
Murthy N S R, Murthy B S. 1962. Natural cross-pollination in chilli. Andhra Agricirltitrul Journal
9: 161-165.
Odland M L, Porter A M. 1941. A study of natural crossing in peppers (Capsicum frutescens).
Proceedings of the American Society for Horticultural Sciences 38:585-588.
Percival M S. 1961. Types of nectar in Angiosperms. New fhytology 60:235-281.
Peterson P A. 1958. Cytoplasmatically inherited male sterility in Capsicmi. American Natitralist
92: 11 1-1 19.
Pouvreau A. 1984. Les plantes a epices: Le piment (Capsicum L., Solanacees); Cultures potageres:
le poivron (Capsicum anniciim L.. Solanacees). In Pollinisation et Production Vkgkrales, pp. 431-
438: 45M52. Eds P Pesson and J Louveaux. Paris: INRA.
Purseglove J W. 1968. Tropical Crops: Dicoyledons I . Dicotyledotis 2. New York: John Wiley &
Sons. 719 pp.
Rylski I. 1986. Pepper (Capsicum). In Handbook of Fruit Set and Development. pp. 341-353. Ed. S
P Monselise. Boca Raton. FL: CRC Press Inc.
Shifriss C, Frankel R. 1969. A new male sterility gene in Capsicurn annicum. Proceedings of the
American Society f o r Horticultural Sciences 94:385-387.
Waller G D. 1972. Evaluating responses of honeybees to sugar solutions using an artificial-flower
feeder. Annals of the Entomological Society of America 65:857-862.
Wykes G R. 1952. The preferences of honeybees for solutions of various sugars which occur in nectar.
Joicrnal o f Experimental Biology 29:511-518.
(Received 28 April 1992)