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Nectar biodiversity: a short review

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DOI: 10.1007/s00606-002-0277-y

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Plant Syst. Evol. 238: 7–21 (2003)
DOI 10.1007/s00606-002-0277-y
Nectar biodiversity: a short review
E. Pacini1, M. Nepi1, and J. L. Vesprini2
1
Department of Environmental Sciences ‘‘G. Sarfatti’’, Siena, Italy
2
Faculty of Agricultural Sciences, Zavalla, Argentina
Received July 23, 2002; accepted September 18, 2002
Published online: June 2, 2003
Ó Springer-Verlag 2003
Abstract. Nectaries differ in many aspects but a
common feature is some kind of advantage for the
plant conferred by foraging of consumers which may
defend the plant from predators in the case of
extrafloral nectaries, or be agents of pollination in
the case of floral nectaries. This minireview is
concerned mainly with floral nectaries and examines
the following characteristics: position in flower;
nectary structure; origin of carbohydrates, amino-
acids and proteins; manner of exposure of nectar; site
of nectar presentation; volume and production of
nectar in time; sexual expression of flower and
nectary morphology; nectar composition and floral
sexual expression; variability of nectar composition;
fate of nectar; energy cost of nectar production. The
species of certain large families, such as Brassicaceae,
Lamiaceae and Asteraceae, resemble each other in
nectary organisation; other families, such as Cucur-
bitaceae and Ranunculaceae, have various types of
organisation. A scheme is presented to illustrate
factors influencing nectary and nectar biodiversity.
Key words: Floral nectaries, nectar, secretion,
carbohydrates.
Introduction
The term nectary does not indicate a well
defined anatomical structure because there are
various types of nectary with different
anatomical origins and positions. The term
has ecological significance, in that nectaries are
the places where liquid substances involved in
interactions with animals are produced and
offered.
The most ancient plant with nectaries still
living today is in Pteridophyta. Pteridium
aquilinum, a quasi-cosmopolitan fern in which
nectaries provide rewards for ants that defend
the plants from predators (Heads and Lawton
1985 and references therein). These authors
demonstrated nectaries in fern populations in
Great Britain, but they were not found in
populations in South Africa (Rashbrook et al.
1992). Thus, ecological significance of nectar-
ies may vary in relation to environment.
In gymnosperms, nectaries are only found
in certain Chlamydospermae such as Ephedra
sp. and Welwitschia, being involved in polli-
nation (Bino et al. 1984, Wetschnig and
Depisch 1999). Nectaries are more common
in angiosperms, dating back to the Late
Cretaceous, when they were both floral and
extrafloral (Friis and Endress 1990). The
former are involved in pollination and the
latter in rewards for animals, mainly ants, that
keep predators away, but in some cases they
are also involved in pollination. Examples of
this type are the cyathial nectaries of the genus
8 E. Pacini et al.: Nectar biodiversity

Euphorbia and the stipules of leaves in Acacia
(Kenrick et al. 1987 and references therein).
Fahn (1952) gives an anatomical and
functional description of floral nectaries, iden-
tifying five types. In a later paper (Fahn 1979),
he identifies 12 topographic positions of floral
and extrafloral nectaries. Vogel (1997, 1998a,
1998b, 1998c) gives an exhaustive description
of the type and structure of nectaries in the
main angiosperm families. Percival (1961)
describes types of nectaries in 889 species of
angiosperms. Nectaries vary in morphology,
nectar varies in chemical composition and
these characteristics reflect the type of pollina-
tor (Baker and Baker 1983).
There are families of angiosperms in which
all species have nectaries, whereas nectaries are
absent in Orchidaceae wherever pollination
occurs by deception (Neiland and Wilcock
1998). These authors noticed that in this
families reproductive success was statistically
greater when nectaries were present.
Schmid (1988) published a paper on termi-
nology which summarises research on nectaries
up to the 1980s. A detailed survey of distribu-
tion, types and evolution of monocot nectaries
was given by Smets et al. (2000); data on dicots
is divided among various papers (Schmid 1988
and references therein).
The present minireview is mainly con-
cerned with floral nectaries. Its aim is to
examine cytological and ecological aspects
rather than anatomical and topographic ones.
The latter are well known and are more easily
studied and interpreted.
Observations and discussion
Position of nectaries in flowers
Floral nectaries may be at the same level as the
surface of the organ that bears them, form a
protrusion, or be deeply embedded, as in the
case of septate nectaries (Fahn 1979). If they

Table 1. Relationships between features of nectary and nectar and some environmental parameters. The
presence of ‘‘X’’ means a correlation between parameters
Pollinator Pollinator Pollen Predation Blooming No. of
type choices charge and period ovules per
discharge ovary
– Floral nectary X X X
position
– Nectary structure X X X
– Nectar carbohydrate X X
source
– Nectar production X X
rate
– Site and mode of X X
nectar presentation
– Nectar volume and X X X X X
production in time
– Flower sexual X X
expression and
nectary morphology
– Nectar composition X X
and flower sexual
expression
– Variability of nectar X X X
composition
– Fate of nectar X X
E. Pacini et al.: Nectar biodiversity
are exposed, as in the case of Hedera helix, the
nectar may be collected by many animals
with different pollination efficiencies. Exposed
nectar also evaporates readily and is not
protected by insect predation. On the other
hand, if the nectar is secreted inside the
corolla, or in a spur, it is protected against
evaporation, especially if the cuticle of the
floral part collecting nectar is thick, but it is
not protected from robbery (Inouye 1983), see
also Table 1.
Nectary structure
The nectary consists of three components: a.
epidermis with or without stomata which
normally mediates nectar release; b. parenchy-
ma that produces or stores substances that
become dissolved in the nectar; c. the vascular
bundle which conveys water or nutrients to the
parenchyma. These three parts may vary
greatly, depending on the type of nectary.
The reasons for this diversity depend largely
on the type of pollinator, the manner in which
pollen is loaded/unloaded and on the number
of ovules per ovary (Table 1).
The epidermis of the nectary may be:
1. flat;
1.1 devoid of stomata as in the case of
Helleborus foetidus (Fig. 1);
1.2 with stomata as in the case of Cucurbita
pepo (Figs. 1 and 2);
2. with secreting hairs as in Cyclanthera pedata
(Figs. 1, 2, 3).
The nectar may be exposed in microdroplets at
the end of hairs or in a continuous layer over
the nectary surface. There is normally a cuticle
(Fig. 4) but it may be perforated to enable
nectar to go out to exit or the cuticle may have
modified permeability, as in C. pepo in which
nectar uncollected by pollinators is resorbed
(Nepi et al. 1996a).
Parenchyma specialised for nectar secretion
may be mono- or multi-layered. Its volume
is proportional to the quantity of nectar
produced and hence the type and number of
pollinator visits to the nectary (Table 1).
9
The vascular bundle which supplies the
parenchyma may consist of the following
elements, depending on secretory mode: a.
phloem only, e.g. floral nectaries of Brassica
napus (Davis et al. 1986), Vicia faba (Davis
et al. 1988), Digitalis purpurea (Gaffal et al.
1998) and Cyclanthera pedata (Fig. 3); b. phlo-
em and xylem: as in extrafloral nectaries of
Vicia faba (Davis et al. 1988) and other
examples in Fahn (2000).
From a systematic viewpoint, there are
families, such as the Brassicaceae, Lamiaceae
and Asteraceae, in which the nectaries are
similar throughout the family (for Asteraceae
see Mani and Saravanan 1999). On the other
hand, at least two different types exist in the
Cucurbitaceae, with nectar being secreted in
two different ways but presented in only one
way (Fig. 1). In Cucurbita pepo, the nectar
comes out of stomata (Figs. 1, 2) and is exposed
in a continuous liquid layer. In Cyclanthera, the
epidermis has multicellular hairs and the nec-
tar, produced in the hair tip, builds up and
forms a continuous layer (Fig. 1).
Origin of nectar carbohydrates
According to Fahn (2000), the origin of
secreted nectar is the phloem sap. Directly or
photosynthesis in the parenchyma. This means
individual and seasonal variations, especially
in plants that bloom over long periods
(Table 1). The parenchyma with chlorophyll
may belong to the nectary itself, as in Cyclan-
thera pedata (Figs. 1, 5a, b), certain Ranuncul-
aceae such as Helleborus (Vesprini et al. 1999),
and Brassicaceae, or it may be in other parts of
the flower or plant, as in Cucurbita pepo (Nepi
et al. 1996b).
In Cucurbita and Passiflora (Table 2),
products of photosynthesis by parts of flowers
other than nectaries or by other parts of the
plant, accumulate in the nectar-producing
parenchyma as starch in amyloplasts, begin-
ning several days before anthesis. Amyloplasts
in the nectar-producing parenchyma are
generally almost devoid of stroma because
they are fully packed with starch (Nepi et al.
10 E. Pacini et al.: Nectar biodiversity

Cucurbita pepo Cyclanthera pedata Helleborus foetidus

1
2
CP
3
Before 4
Nectar 5
secretion secretion
AP 6 CP
AP CP
AP 7
CP
8

1
2
CP
3
4
Nectar 5
exposure PP 6
PP PP 7 CP
CP CP 8

secretion
Nectar

During
consumption 1
2
3 CP

Eventual 4
nectar 5
resorption AP 6
CP
AP 7
AP
8

Fig. 1. Scheme of formation, exposure and presentation of nectar in three species differing in aspects such as
origin of sugar component of nectar, presentation and reabsorption. Nectar is shown as dots. Amyloplasts
(AP), proplastids (PP), chloroplasts (CP). In Cucurbita pepo nectar is derived from hydrolysis of starch in the
parenchyma. Nectar is exposed as a continuous layer on the epidermis and is resorbed if not consumed. In
Cyclanthera pedata it is produced by photosynthesis by the parenchyma, secreted through pluricellular
trichomes and presented as a continuous layer on the epidermis. If not collected, it is not resorbed. In Helleborus
foetidus the nectary surface is disposed vertically, the sugar component is produced by the parenchyma but the
nectar is exposed gradually by epidermal cells which empty completely and die. After daily secretion, the
epidermis and the nectary reorganise. The nectar flows along the nectary and that not collected falls to the
ground

c
Fig. 2. Cucurbita pepo PAS stained nectary producing nectar (arrows). There is a monolayered epidermis (E )
with stomata (s) and nectar producing parenchyma (P) containing starch. Nectar accumulates in the stomata
cavities (arrow) and on the surface of the nectary (arrowheads). Nectar is PAS positive, viscous and not
removed during fixation and dehydration. Bar ¼ 50 lm. Fig. 3. Cyclanthera pedata PAS stained nectary. The
nectary is of the trichome type consisting of a monolayered epidermis (E ) with secretory capitate trichomes and
parenchyma with chloroplasts, not evident in the picture, with vascular bundles (vb) and phloem. The trichomes
have a peduncle consisting of two cells and an apex consisting of 6–7 cells. Bar ¼ 50 lm. Fig. 4. Cyclanthera
pedata auramine O stained nectary. A cuticle (arrow) is more evident above the trichome than above the other
epidermal cells. Bar ¼ 75 lm. Fig. 5a and b. Hand section of Cyclanthera pedata nectary observed in bright
field (5a) and UV light (5b); fluorescence is from chloroplasts in parenchyma cells. Trichomes do not contain
chloroplasts and do not fluoresce. Bar ¼ 200 lm
E. Pacini et al.: Nectar biodiversity 11
12
1996b). Moreover, there are many starch
granules per amyloplast; this increases starch
surface area, facilitating and speeding hydro-
lysis at the time of nectar production and
secretion. The starch is hydrolysed before the
start of secretion (Figs. 1, 2) as seen in
Cucurbita (Nepi et al. 1996a, b), Passiflora
sp. (Durkee et al. 1981), Hibiscus rosa-sinensis
(Sawidis 1998), Rosmarinus officinalis (Zer and
Fahn 1992), Pisum sativum (Razem and Davis
1999) and the orchid Limodorum abortivum
(Figueiredo and Pais 1992). Nectar production
may vary in quantity (Table 2); when it is very
abundant amyloplasts may even accumulate in
epidermal cells (Nepi et al. 1996b). The abun-
dance of starch in parenchyma cells of Cucur-
bita pepo and Passiflora biflora (Nepi et al.
1996b, Durkee 1982, Durkee et al. 1981) has
been linked to the high rate of production of
nectar with a high sugar concentration.
Depending on the origin of the nectar, the
nectary parenchyma may be of two kinds, it
may have chlorophyll or it may be reserve
parenchyma for starch storage (Figs. 1, 6). In
the former, starch is produced directly by
photosynthesis and occurs in small quantities,
stored for only a few hours, to be hydrolysed
at night; photosynthesis may proceed for many
days, producing small quantities of starch and
nectar every day.
There would seem no reason why both
types of parenchyma should not occur simul-
taneously or in succession, as reported in
Sanango racemosum (Gesneriaceae) (Maldona-
do and Otegui 1997). In this species the nectar-
producing parenchyma also contains starch.
According to these authors, the starch does not
contribute to nectar production but is, never-
theless, gone by the end of anthesis.
Such a process would be regulated by
plastid conversion processes, as amyloplasts
are normally a final stage of plastid differen-
tiation (Wheatley 1977).
Once nectar secretion has ended, the fate of
the two types of parenchyma is again different.
Photosynthesizing parenchyma may continue
to function for a while, with the products
going to other activities, such as growth of the
E. Pacini et al.: Nectar biodiversity
ovary. Finally, the nectary degenerates. Stor-
age parenchyma just seems to degenerate as in
Cucurbita (Nepi et al. 1996b).
In Helleborus bocconei and H. foetidus both
of which have both types of parenchyma,
the two parenchymas have the same mode
of secretion but the origin of the sugar compo-
nent of nectar is slightly different. In H. bocc-
onei, the photosynthesizing parenchyma seems
to produce all the nectar, whereas in H. foetidus
there must be a contribution from photosyn-
thesis by other parts of the flower and/or plant.
H. bocconei has an intense green and exposed
nectary and its cells are packed with chlorop-
lasts; the nectar, predominantly sucrose (mean
sucrose concentration 275.18 mg/ml), is pro-
duced when the flower is exposed. In H. foetidus
the nectary is always protected within the
flower, it is pale green and the nectar, which
is sucrose dominant (607.81 mg/ml), is pro-
duced before the flower opens. These characters
and the different mean sucrose contents suggest
that some of the nectar originates elsewhere
(Vesprini et al. 1999 and our unpublished data).
The two modes of nectar carbohydrate
production may serve to produce nectar con-
sumed by pollinators at different times of day
(Fig. 6). If sugars are derived from photosyn-
thesis in the parenchyma itself, they may be
produced over a long period, even weeks as in
the floral nectaries of Eucalyptus (Davis 1997)
and Helleborus (Vesprini et al. 1999), and in all
extrafloral nectaries (Table 2). However,
because production only occurs by day, the
nectar must be ready at dusk if pollination is
nocturnal. To have nectar ready at sunset,
starch stored in chloroplasts during the day
would have to be quickly hydrolysed at dusk.
That is unlike what happens in leaves, where
hydrolysis is slower and continues throughout
the night. Photosynthesizing nectaries have
intermittent nectar production because nectar
can be produced only during the day. However
if material for nectar sugar comes from
hydrolysis of starch stored in the parenchyma,
nectar may be produced at any time of day,
even before sunrise. By this method it is also
possible to produce much nectar in a short time.
E. Pacini et al.: Nectar biodiversity 13

Table 2. Presence and abundance of starch in nectary parenchyma, duration of secretion in floral (F ) and
extrafloral (E ) nectaries of various species. When stored starch is abundant, it is even deposited in the
epidermis (ep). Extrafloral nectaries never store starch and are of the photosynthetic type. Floral nectaries
produce nectar by photosynthesis of the nectary or other parts of the flower/plant. Abundance of
amyloplasts in nectary parenchyma is reported by one or more +. Chloroplast presence in the nectary
parenchyma is reported by +
Species and Nectary Plastid type Duration of Author
family type secretion
Amyloplasts Chloroplasts
Brassica napus F + Davis et al. 1986
(Brassicaceae)
Citrus sinensis F + Rachmilevitz and
(Rutaceae) Fahn, 1973
Cucurbita pepo F +++(ep) several hours Nepi et al. 1996
(Cucurbitaceae)
Deuterochonia E + several days/ Galetto and
longipetala some weeks Bernardello 1991
(Bromeliaceae)
Dyckia floribunda E + several days/ Galetto and
(Bromeliaceae) some weeks Bernardello 1991
Euphorbia nerifolia E + several day/ Arumugasamy et al.
(Euphorbiaceae) some weeks 1990
Limodorum F + Figueiredo and
abortivum Pais 1992
(Orchidaceae)
Lonicera F + Fahn 1979
japonica
(Caprifoliaceae)
Musa paradisiaca F + Fahn and Benouaiche
(Musaceae) 1979
Passiflora biflora F +++(ep) several hours Durkee et al. 1981
(Passifloraceae)
Ricinus communis E + several days/ Baker et al. 1978
(Euphorbiaceae) some weeks
Rosmarinus F ++ (ep) Zer and Fahn 1992
officinalis
(Lamiaceae)
Sambucus nigra E + several days/ Fahn 1987
(Caprifoliaceae) some weeks
Vicia faba F + Davis et al. 1988
(Fabaceae)
Vinca rosea F + Rachmilevitz and
(Apocynaceae) Fahn 1973

The difference between the two modes of nectar starch, however no census has been made.
carbohydrate formation are compared in Fig. 6. Table 2 compares nectary type with mode of
In our experience, photosynthesizing nec- nectar production. Extrafloral nectaries are
taries are more common than those that store always of the photosynthesizing type.
14 E. Pacini et al.: Nectar biodiversity

of the nectary nectar is produced for a few • low rate of secretion

hours or several days • the nectar produced is proportional
to mass and volume of nectary
• the pollinator requires little nectar
• the nectary stops secreting nectar
but photosynthesis continues and its
products may be used for other
purposes

Examples: Brassicaceae,
Lamiaceae,
Chloroplasts
Fabaceae
some Scrophulariaceae

of other parts of products of photosynthesis starch is hydrolyzed and • high rate of secretion
the flower or are stored in the nectary nectar is available at any • large amounts of nectar can be
leaves/stem parenchyma time of day produced in a short time
• pollinators consume large amounts
of nectar
• the nectary degenerates

Examples:
Cucurbita pepo
some Malvaceae

Fig. 6. Scheme showing the two possible origins of the carbohydrate part of nectar and eco-physiological
consequences

Origin of the amino acid and protein cytoplasm enrich the nectar with proteins and
components of nectar amino acids (Fig. 1). The emptying of epider-
mal cells as they secretes causes changes in
There has been even less research on the origin
position of all epidermal cells (Fig. 1); those
of the amino acid and protein components of
that have secreted nectar are displaced and
nectar than on the carbohydrate components.
deformed by the growth of those about to
It is likely that the site of production of amino
secrete.
acids in nectar is not always the same and that
it varies in relation to nectary structure. This
makes it interesting to discover the relation Manner of exposure of nectar
between nectar composition and nectary struc-
The manner in which nectar is produced and
ture.
exposed is summarised in Kronestedt-Ro-
A useful case is that of Helleborus, in which
bards and Robards (1991). Here, we include
there is some certainty as to where all or most
their data together with that of more recent
of the amino acid component is produced. The
studies on nectar exposure. Nectar may
epidermal cells of the nectary of Helleborus die
appear on the surface of the nectary in four
as they produce nectar (Vesprini et al. 1999).
ways.
Sugars derived from the underlying photosyn-
thesizing parenchyma pass through the epider- 1. It comes out of modified stomata which
mal layer and proteins from the degenerating have lost the capacity to open and close,
E. Pacini et al.: Nectar biodiversity
as in Vicia (Davis and Gunning 1992),
Cucurbita (Nepi et al. 1996b) (Figs. 1, 2)
and Linaria (Nepi et al., this volume).
2. It crosses the plasma membrane then the
cell wall and accumulates in a pocket
between the cell wall and cuticle. The cuticle
breaks under the nectar pressure. This
occurs in Anacardium occidentale (Wunna-
chit et al. 1992).
3. The cell wall has ingrowths like transfer
cells that facilitate and speed secretion as in
Lonicera japonica (Fahn 1979). This mode is
found mainly in pedunculate nectaries such
as those of Abutilon (Kronestedt et al.
1986).
4. The nectar flows out through the external
periphery of the epidermis which has a
cuticle, as in Limodorum abortivum spur
(Figuereido and Pais 1992).
5. The epidermal cells gradually die, producing
nectar for a given period (Fig. 1) as in
Helleborus (Vesprini et al. 1999).
Clearly the way nectar is exposed and
presented determines the type of nectar col-
lector and robbers (Table 1). Case 1 is perhaps
the most common. All parenchyma and epi-
dermal cells produce and secrete nectar at the
same time. In cases 2, 3 and 4, parenchyma
cells operate simultaneously and secretory
epidermal cells are synchronous in the pro-
duction/exposure of nectar. In case 5, the
secretory epidermal cells are asynchronous
(Fig. 1).
Site and mode of nectar presentation
to pollinators
Just as pollen has primary and secondary
presentation (Faegri and van der Pijl 1979), we
propose the same for nectar. Primary presen-
tation is exemplified when nectar is exposed to
pollinators on: a. the surface of the nectary
itself, like pollen presented in the anther; b. on
continuous surfaces; c. or on the surface of
capitate nectaries (Fig. 1). Presentation is
secondary if the nectar flows from the nectary
and collects in another part of the flower, as in
15
spurs or similar structures the task of which
seems to be to hold the nectar until it is
collected. Obviously the position, length and
accessibility of the spur determine the type of
collector and any predators. The spur may be
the nectary itself if it produces nectar as in
Limodorum abortivum (Figueiredo and Pais
1992) or it may merely be where nectar
accumulates if the nectary is in another part
of the flower, as in Linaria (Nepi et al., this
volume) and the spur is a recipient. In
Asclepias syriaca the nectary is in the stigmatic
chamber and the nectar flows from there in
nectar reservoirs (cuculli) (Kevan et al. 1989).
There is an intermediate case in vertically
disposed bell-shaped or tubular flowers and
nectaries. In some Aloe species the nectary is at
the base of the corolla and the nectar flows
along the corolla tube; in Helleborus, the
nectary performs photosynthesis and is tubular
(Vesprini et al. 1999); the nectar flows along
the nectary itself (Fig. 1).
Volume and production of nectar in time
The volume and production of nectar in time
are influenced by environmental parameters
(Table 1). The quantity of nectar produced by
a nectary varies enormously from fractions of
a microlitre (difficult to detect) to thousands
of microlitres (Opler 1983). Abundant pro-
duction of nectar may be related to numerous
pollinator visits (Waddington 1983), especial-
ly in species with many ovules/ovary, or to
large pollinators. The larger the pollinator,
the more the nectar consumed per visit, the
larger the body surface for collecting and
depositing pollen. Repeated visits create suc-
cessive waves of pollen tubes in the style
(Nepi and Pacini 2001), reducing male and
female competition and enabling scalar fertil-
isation of ovules in the ovary, especially in
species with many ovules (Nepi and Pacini
2001).
The nectar may be produced for a brief
time (only a few minutes or hours) or for
many days. When much nectar is produced in
a short time, it normally comes from hydro-
16
lysis of starch stored in the parenchyma
(Fig. 6).
Sexual expression of flowers
and nectary morphology
In the case of unisexual flowers, nectaries may
be on both flowers or on only one, as in Osyris
alba (Aronne et al. 1993) where they are on
male flowers and pollinators are induced to
visit female flowers by deception (Table 1). A
similar situation is found in Ecballium elateri-
um where nectaries of male flowers produce
nectar and those of female flowers do not
secrete measurable amounts (Fahn and Shimony
2001). In Cucurbita pepo, flowers of both sexes
have nectaries but they are dimorphic in posi-
tion and form, causing different patterns of
nectar collection which optimises pollen collec-
tion and deposition (Nepi and Pacini 1993).
Moreover, the nectar of male and female flowers
differs in quantity and composition in Cucurbita
pepo and Cocos nucifera (Kevan 1993; Nepi et
al. 1996a, 2001). The nectaries of male and
female flowers are also different in Croton
sarcopetalus (Freitas et al. 2001) and female
flowers have two types. Septate nectaries of
Euterpe precatoria are only slightly different
(Küchmeister et al. 1997).
Sexual expression of flowers
and nectar composition
Nectar composition may differ in male and
female flowers of monoecious and dioecious
species. That produced by female flowers of
Cucurbita pepo, Euterpe precatoria and Rhus
hirta is more abundant and more concentrat-
ed than that of male flowers (Greco et al.
1996, Küchmeister et al. 1997, Nepi et al.
2001). On the other hand, in the three species
of the genus Silene, Witt et al. (1999) found
more concentrated nectar in male flowers. In
Cucurbita pepo, it has been observed that the
difference in quality and quantity of nectar in
the two flowers is reflected by the fact that
female flowers receive more visits (Table 2)
(Nepi and Pacini 1993; Nepi et al. 1996a, b).
E. Pacini et al.: Nectar biodiversity
In hermaphrodite flowers, nectar composition
may vary in relation to the sexual stage of the
flower. For example in Alstroemeria aurea
and Echium vulgare, nectar produced in the
male phase is more abundant and concentrat-
ed (Klinkhamer and de Jong 1990, Aizen and
Basilio 1998), whereas in Polyascias sambuci-
folia that produced in the female phase is
more concentrated (Gillespie and Henwood
1994).
Variability of nectar composition
Besides programmed variability related to
sexual expression of the flower, there are other
types of variability that depend on the envi-
ronment and the plant itself (Table 1).
There is variability in flowers of the same
plant, or of flowers of individual plants of
population of the same species, or between
different populations of the same species.
These different types of variability probably
at least partly reflect different photosynthetic
activity of the plants or of various parts of a
plant, especially in the case of trees. Because
nectar is derived directly or indirectly from
photosynthesis, its composition may vary
widely in relation to exposure of the flower,
or parts of the plant carrying flowers, to light,
and consequently in relation to temperature,
and availability of water. This explains why
temperature can affect nectar secretion and
composition (Jacobsen and Kistjánsson 1994).
Freeman and Head (1990) demonstrated that a
positive correlation exists between increase in
temperature, obviously within limits, and in-
crease in nectar concentration. If nectar pro-
duction and composition are influenced by
temperature, it follows that nectar quality
would change in time in species with long
blooming times, such as Capparis spinosa
(Petanidou et al. 1996) as well as with the age
of the plant, as in Silene dioica (Hemborg
1998). Pleasants and Chaplin (1983) demon-
strated in Asclepias quadrifolia a positive
correlation between nectar production and
root weight, and a negative one between nectar
production and inflorescence size.
E. Pacini et al.: Nectar biodiversity 17

Fate of nectar and nectary of nectaries can be up to 37% of daily

photosynthesis (Southwick 1984, Pyke 1991).
Nectar produced by a nectary may be: a. stolen
This may be why plants that produce abun-
by animals that do not pollinate; b. collected
dant nectar resorb what is left over (Nicolson
wholly or partly by animals that pollinate;
1995, Nepi et al. 2001, Nepi et al. this
c. left for lack of pollinators (Table 1). Nectar
volume).
not consumed may fall out of the flower or
There is also another case of nectar saving.
nectary, especially if they are tubular and hang
Cleistogamy is a temporary phenomenon in
down (Fig. 1), to the benefit of soil microor-
many plants which have flowers that do not
ganisms. In some cases it is resorbed by the
open and automatically self-pollinate. Cleisto-
plant. The best known examples of nectar
gamic flowers are always smaller than those
resorption are documented in Cucurbita, Lina-
capable of cross pollination (Lord 1981). The
ria (Nepi et al. 2001, Nepi et al. this volume),
advantage of cleistogamy is that it produces
Grevillea (Nicolson 1995) and Eucalyptus
seeds with less energy investment, albeit with
(Davis 1997) which differ in aspects such as
reduced variability. Some species subject to
different origin of nectar carbohydrates, dura-
temporary cleistogamy are anemophilous,
tion of anthesis, site of nectar exposure, and so
though there are many entomophilous (Lord
forth. The corolla closes at the end of anthesis
1981), some of which have nectaries. At least in
in Cucurbita and in Linaria it is always closed
some species of the genus Viola (V. canina,
by the labium. In Grevillea and Eucalyptus,
V. hirta and V. odorata) cleistogamic flowers
nectar droplets remain unprotected during
are devoid of nectaries (our unpublished data).
resorption (Nicolson 1995).
However, we do not know to what extent this
When nectar secretion ceases, the fate of
feature occurs among species manifesting cleis-
the nectary tissue may depends on its position
togamy.
in the flower and on the origin of the carbo-
The lack of nectaries in cleistogamic flow-
hydrate part of the nectar. In fact when nectar
ers of the genus Viola is an example of
sugars derive from starch hydrolysis, the nec-
coordination of plant activity towards energy
tary falls or degenerates because it can not play
saving. The case of Pittosporum tobira is
any other function.
different: this plant is male sterile meaning
When nectar sugars derive from the pho-
that the anthers do not produce pollen, but it is
tosynthetic activity of the nectary, it may fall,
apomictic because fruits ripen, dispersing seeds
as in the case of Brassicaceae, or persist. It
which germinate. Despite this, the plant has
persists when the nectary is a part of the ovary
nectaries and bees visit the flowers and collect
either under (Linaria vulgaris, Ericaceae etc.)
nectar (Ricciardelli D’Arbore and Persano
or on the ovary (Hedera helix, Brassicaceae,
Oddo 1978), to no apparent benefit of the
some Aloe species). In this case the photosyn-
plant.
thetic activity of the nectary may continue and
the product may contribute to fruit develop-
ment. This is only a reasonable hypothesis and Future research
researches are necessary to confirm it.
Existing knowledge on nectaries can be defined
as abundant but not integrated or rationalised.
Cytological aspects have been related rarely to
Energy cost of nectar
physiological and ecological aspects. Apart
Histologically, nectaries consist of secretory from reviews, papers presenting the results of
tissue and nectar is exposed to reward research usually regard a single species, which
seeking pollinators or animals that defend means comparisons of different types of nec-
the plant. Nectar is therefore an investment tary is rare. Certain cytological and ecological
that may be costly in energy terms. The cost aspects have never been studied (e.g. the
18
presence of cuticle and the nectariferous
parenchyma type) and because laboratory
and field study techniques are not standar-
dised, certain data cannot be compared
between species.
Apart from the fundamental contribution
of Bonnier (1879) and Vogel (1997, 1998a, b,
c), there is no systematic view of the various
cytological and ecological aspects of nectaries
in angiosperm families.
In our opinion, there are still gaps in our
knowledge of the origin of the polysaccharide
component of nectar. Only a few reports
demonstrate an effect of environmental and
experimental conditions on nectar composition
and abundance (Corbet et al. 1979, Wyatt et al.
1992). Vickery and Sutherland (1994) analysed
the volume and concentration of nectar in
spontaneous and cultivated plants and found
that concentration changes little whereas vol-
ume changes considerably, especially in green-
house plants.
There have been various studies on nectar
composition, mostly on soluble monosaccha-
rides and oligosaccharides, but not on the
presence of PAS-positive polysaccharides that
survive fixation, like those observed in Cucur-
bita pepo (Fig. 2). No explanation has been
offered for why nectar, a sugar solution, is not
removed by fixation and dehydration and also
shows PAS positivity indicative of polysaccha-
rides. All this suggests that nectar contains
unknown dissolved polysaccharides.
Little work has been done into differences
between nectaries and nectar composition in
plants with wide distributions, for example the
ecological studies on Pteridium aquilinum
(Heads and Lawton 1985, Raschbrook et al.
1992).
Although it is fairly clear where the sugar
component of nectar comes from, the origin of
the amino acid-protein and other components
has received less attention. Research on this
topic would be useful.
It would also be useful to know if our
observations on the lack of nectaries in cleis-
togamic flowers of Viola are a feature of all
cleistogamic plants with nectaries. Other
E. Pacini et al.: Nectar biodiversity
aspects that deserve attention are the relations
between nectary types and breeding systems
e.g. dioecy, monoecy, autogamy, xenogamy
and parthenocarpy.
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21
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Addresses of the authors: E. Pacini (e-mail:
pacini@unisi.it), M. Nepi, Department of Envi-
ronmental Sciences ‘‘G. Sarfatti’’, Via P. A. Matt-
ioli 4, I-53100 Siena, Italy. J. L. Versprini, Faculty
of Agricultural Sciences, UNR CC14 2123 Zavalla,
Argentina.