Professional Documents
Culture Documents
1
Objectives
o To determine the pollination syndromes of selected 10 flowers by looking at their
floral morphology
o To infer the breeding systems of selected 10 flowers by using pollen: ovule ratio
2
Introduction
Plant breeding systems have been viewed as mechanisms to promote outcrossing (or as
mechanisms to prevent inbreeding). As mentioned above, hermaphrodites are common in plants,
while much fewer species are dioecious. Dioecy is easily seen as a mechanism to prevent
inbreeding, since male and female flowers exist on separate plants. Hermaphrodites, on the other
hand, must evolve other mechanisms to prevent selfing, or at least, to reduce inbreeding
depression (i.e. the accumulation of deleterious, recessive alleles). Below is a list of some types
of breeding systems:
Dichogamy refers to temporal separation in male and female functions of bisexual (or
perfect flowers). Protandry occurs when pollen sheds first, before the stigmas become receptive.
Protogyny, on the other hand, occurs when the stigmas become receptive first, before pollen is
released. Protandry is more common than protogyny. In order for dichogamy to be an effective
outcrossing mechanism, all flowers on the plant must be synchronous (i.e. if protandry occurs, all
flowers must release their pollen before the stigmas become receptive). Otherwise, pollen can be
transfered from one flower to another on the same plant (this is referred to as “geitonogamy”).
3
Dichogamy can also work if there is a sequential developmental pattern of male and female
flowers (i.e. certain pollinators tend to move ‘up’ a plant - if flowers are male first, and develop
into female flowers starting from the bottom of the plant, then the chances of geitonogamy can
be reduced).
Herkogamy is the spatial separation of styles and anthers in a perfect flower. This system
is apparently widespread; however, geitonogamous pollen transfer is possible. In flowers that do
not have spatial separation of styles and anthers, self pollination can occur.
Sex-switching simply refers to a plant that is either male or female at one time, and then
later (perhaps later in the growing season), the plant switches to the opposite sex.
4
there is a parental effect with the sporophytic system). Gametophytic SI is found in Oenothera
sp., as well as in the Solanaceae (potato family), Papaveraceae (poppy family), Poaceae (grass
family), Ranunculaceae (buttercup family), and the Rosaceae (rose family, e.g. apples).
Sporophytic self-incompatibility is found in the Brassicaceae (mustard family), Asteraceae (aster
family), and the Convolvulaceae. It had been oringially thought that self-incompatibility systems
were responsible for the success of the angiosperms (i.e. self-incompatibility was considered to
be an ancestral trait in the angiosperms). By comparing these different systems, it is now known
that self-incompatibility arose independently on numerous occassions (e.g. some genes involved
in these systems have been sequenced and compared -- these genes encode very different
products in different species).
Flower is the sexual reproductive structure of the Angiosperms, adapted for pollination.
The flower consists of a determinate, modified shoot (the floral axis or receptacle) bearing
modified leaves (the perianth parts, stamens and/or carpels). There are two types of flowers; an
‘imperfect’ flower has either male or female components, but not both. A ‘perfect’ flower has
both male and female components. A perfect flower is also known as a hermaphrodite or as a
‘bisexual’ flower.
Pollination is the deposition of mature pollen/s on receptive stigma of the same flower or
some other flower of the same species. In pollination the male and female gametes are mixed to
produce the offspring. According to pollination type the angiosperms are of two categories;
Allogamous plants and Autogamous plants. Allogamous plants highly rely on out breeding
5
(cross pollination) with other individuals of the species, whereas Autogamous plants rely on
selfing (self pollination).
The allogamous plants have to transfer the pollens from one plant to another. Therefore
they should employ many pollen transfer systems. These pollen transfer systems can be
commonly referred to as pollination syndromes.
Pollination syndromes
Flowers may be small and inconspicuous, green and not showy. They produce enormous
numbers of tiny pollen grains (hence wind-pollinated plants may be allergens, but seldom are
animal-pollinated plants allergenic). They have large feathery stigmas to catch the pollen grains.
They grow in low-diversity stands and are among the taller species in their communities. Insects
may visit them to collect pollen, but they are not the most effective pollinators and exert little
selection pressure on them.
Water-pollinated plants are aquatic. Their flowers tend to be small and inconspicuous with lots
of pollen grains and large, feathery stigmas to catch the pollen. Many aquatic plants are insect-
pollinated, with flowers that emerge into the air.
Beetle-pollinated flowers are usually large, greenish or off-white in color and heavily
scented. Scents may be spicy, fruity, or similar to decaying organic material. Most beetle-
pollinated flowers are flattened or dish shaped, with pollen easily accessible, although they may
6
include traps to keep the beetle longer. The plant's ovaries are usually well protected from the
biting mouthparts of their pollinators. Beetles may be particularly important in semi-desert areas,
like South Africa and southern California.
There are two types of fly pollination: myophily and sapromyophily. A diversity of flies
(particularly bee flies (Bombyliidae), hoverflies (Syrphidae), etc.) feed on nectar and pollen as
adults, and regularly visit flowers, while male fruit flies (Tephritidae) are attracted to and feed on
specific floral attractant, which acts as fly's sex pheromone precursor or booster, of some wild
orchids (Bulbophyllum species - with highly moveable lip) that do not produce nectar. These are
the myophiles. Sapromyophiles, on the other hand, normally visit dead animals or dung. They
are attracted to flowers that mimic these odoriferous items. They obtain no reward and would
quickly leave, but the plant may have traps to slow them down. These plants have a strong,
unpleasant odor, and are brown or orange in color. They are not as common as myophilous
plants. Myophilous plants do not tend to have a strong scent, and tend to be purple, violet, blue,
and white, open dishes, or tubes. Flies generally utilize many different sources of food making
their pollinating activity infrequent and unreliable. However, their sheer numbers and the
presence of some flies throughout the year make them important pollinators for many plants.
1. Showy, open, bowl-shaped flowers that are relatively unspecialized (e.g. wild roses,
sunflowers)
2. Showy, complicated, non-radially symmetric flowers that are more specialized (e.g.
peas, foxgloves)
7
Some bee flowers tend to be yellow or blue, often with ultraviolet nectar guides and scent.
Nectar, pollen, or both are offered as rewards in varying amounts. The sugar in the nectar tends
to be sucrose-dominated.
There are diverse types of bees, however. Honeybees, bumblebees, orchid bees, etc are
large groups that are quite distinctive in size, tongue length and behavior (some solitary, some
colonial). Thus generalization about bees is difficult. Some plants can only be pollinated by bees
because their anthers release pollen internally, and it must be shaken out by buzz pollination.
Bees are the only animals that perform this behavior.
Bee pollination from mobile beehives is of great economic value for orchards such as
apple or almond.
The plants that are pollinated by moth (nocturnal butterflies) are called sphingophilous or
phalaenophylous plants. These plants have large nocturnal white flowers whit a strong sweet
scent. These features allow moths that are active at night to find the flowers from afar. The moth
stay on the wing while inserting their long tongue into the deep nectar container, These plant
comprise the longest nectar bearing flowering tubes known in the plant kingdom.
Although hummingbirds are the most familiar nectar-feeding birds for North Americans,
there are analogous species in other parts of the world: sunbirds, honeyeaters, flowerpeckers,
8
honeycreepers, bananaquits, flowerpiercers, lories and lorikeets. Hummingbirds are the oldest
group, with the greatest degree of specialization on nectar. Flowers attractive to hummingbirds
that can hover in front of the flower tend to be large red or orange tubes with a lot of dilute
nectar, secreted during the day. Since birds do not have a strong response to scent, they tend to
be odorless. Perching birds need a substantial landing platform, so sunbirds, honeyeaters, and the
like are less associated with tubular flowers.
Bat-pollinated flowers tend to be large and showy, white or light coloured, open at night
and have strong odours. They are often large and bell-shaped. Bats drink the nectar, and these
plants typically offer nectar for extended periods of time. Sight, smell, and echo-location are
used to initially find the flowers, and excellent spatial memory is used to visit them repeatedly.
In fact bats can identify nectar-producing flowers using echolocation, a talent that was only
recently discovered. In the New World, bat pollinated flowers often have sulfur-scented
compounds, but this does not carry to other parts of the world. Bat-pollinated plants have bigger
pollen than their relatives.
9
Determination of pollination syndromes using floral characters
10
Visitation
Mode of Flower
& Color
Pollination Morphology Odor
Anthesis
Carrion & Dung Day & Actinomorphic, Purple, brown Strong, like rotting
Beetles & Night deep corolla tube (like meat, really awful
Flies with appendages meat), no
forming traps nectar
guides or
nectar
Visitation
Flower
Mode of & Color
Morphology Odor
Pollination Anthesis
OCI values;
0 = Cleistogamy
1 = Obligate autogamy
2 = Facultative autogamy
12
Pollen: Ovule ratio, as a method of estimating the breeding
systems
Use of pollen ovule ratio as a method of estimating the breeding system was developed
by Cruden R.W. (1977)
2.4-5.4 Cleistogamy
13
Determination of pollination syndromes using floral characters
“Harmonic” relations between pollinator and perianth shape and color (greatly simplified,
adapted from faegri and van der Pijl,1979).
syrphids
birds
14
Materials and method
Determination of pollination syndrome using floral
morphological characters
Flower shape, corolla type, petal color, flower/inflorescence size, reward type and
amount, timing of the ten flowers were determined.
Ripe anther was selected from each of the flowers of the each species and was
dissolved in distilled water (0.5 mL). One drop (0.05 mL) from the above prepared suspension
was placed on a microscopic slide and the number of pollens was counted under light
microscope. Two replicates were done for each flower.
o Ovule counting
Number of ovules in each flower was counted by observing cross sections of ovaries
under dissecting microscope. Two replicates were done for each flower.
15
Results and Calculations
Determination of pollination syndrome by using floral
morphological characters
16
Traits of flower
Reward Pollination
Name of the flower
Shape Size Colour type & Timing syndrome
amount
Clitoria ternatea Papilionate Small Blue Floral Day Ants
(Katarolu) Zygomorphic tissues,
No scent
and nectar
Tecoma stans Bell shape medium Yellow Strong Day Birds, ants
(Kelanitissa) Actinomorphic plescent
odor,
Some
amount of
watery
nectar
Gardenia Actinomorphic Large White Strong Day Ants, beetles,
grandiflora colour pleasant flies, bees
(Gardenia) Odor,
Nectar
17
present
Determination of breeding systems using out crossing index
(Cruden, 1977)
18
Cerbera manghas a) 52 mm 3 Self compatible,
(Kaduru) b) homogyny 0 3 some demand for
c) same level 0 pollinators
Clitoria ternatea
No. of anthers per flower = (9+9)/2
=9
No. of pollens per drop (0.05 ml) of suspension = (164 +146)/2
= 155
No. of pollens per 0.5 ml of suspension = 155/0.05*0.5
(No. of pollens per anther) = 1550
No. of pollens per flower = 1550*9
= 13950
No. of ovules per flower = (5+5)/2
=5
Name of the flower No. of No. of No. of pollens No. of Pollen: Breeding
anthers per pollens per flower ovules Ovule system
flower per anther per ratio
flower
Clitoria ternatea 9 1550 13950 5 2790 Obligate
(Katarolu) xenogamy
19
Cassia fistula Obligate
(Ehela) 7 42400 296800 60 4946.67 xenogamy
Gardenia Obligate
grandiflora 7 2120 14840 1 14840 xenogamy
(Gardenia)
20
“Harmonic” relation between pollinators and perianth shape
and colour
Nerium oleander Trumpet shaped (f, a) Pink, white, purple Moths, butterflies
(Kaneru)
Mirabilis jalapa Trumpet shaped (f, a) Bright purple, pink, Butterflies, moths
(Hendirikka) yellow, white
Camellia sinensis Dish shape (Unf, a) White, yellow Beetles, flies, bees
(Tea)
21
(Unf, corolla unfused; f, corolla fused; a, corolla actinomorphic; z, corollo zygomorphic)
Discussion
Floral morphological characters play an important role in the determination of pollination
syndrome of a particular flower, because flower shape, color, type of reward, amount of reward
and timing are the deciding factors of the pollination syndrome which is employed by a
particular flower.
Wind-pollinated flowers In general are green, small, and often lack petals. The anthers
and stigmas generally hang outside the flowers to allow the wind to carry the pollen.
As insects, bees are relatively intelligent and are able to learn how to locate and operate
particular species of flowers that are in bloom at a particular time. They are also relatively
strong and are able to push their way into complicated flowers that are not accessible to other
insects.
Bee vision is most sensitive toward the violet end of the spectrum. Therefore, bee-
pollinated flowers tend to have blue or violet markings. Some may even have markings that are
only apparent in the UV range.
Bee-pollinated flowers often have a lobe that serves as a landing pad. Anthers often are
located at the top of tubular petals, dusting the back of the bee as it enters.
22
Plants employ a number of methods to make their flowers more noticeable to pollinators.
All members of the large and successful Asteraceae (sunflower) family have composite
inflorescences containing many flowers arranged to draw attention to the display.
Some flowers produce a bad smell and have a purplish color to simulate the rotting flesh
of dead animals. These flowers attract beetles and carrion flies that pollinate the plant as they are
fooled into trying to lay eggs on the flower. This group includes the Araceae family, which
produces highly modified flowers.
Many hummingbird pollinated flowers are red, a color to which bird eyes are sensitive,
but which is not as apparent to insects. The hummingbird must hover and reach deep inside the
flower to reach the reward. Different species of flower may dust the bird at different locations
on its body so that its pollen will be more likely to end up on another species of the same kind.
These flowers usually do not have a strong odor because the hummingbirds do not have a
particularly well-developed sense of smell.
Moth-pollinated flowers are usually white and have a strong scent. These features allow
moths that are active at night to find the flowers.
The experiment was done to determine the pollination syndrome and breeding system of
ten selected flowers. Four methodologies were being used; Determination of pollination
syndromes using floral characters, Determination of breeding systems using out crossing index,
Estimation of breeding system by using pollen ovule ratio (P:O ratio) and by using Harmonic
relation between pollinators and perianth shape and color.
When looking at floral morphology, there may be several problems which can arise.
• The flower shape can not be exactly determined, as it depends on the
observations. Also there can be more or less similar floral shapes.
• In the case of determining odor, some can be determined an odor as pleasant,
some can determine it as unpleasant, as it is subjective.
• The nectar composition can not be simply determined.
• Time can be either pollinator visitation time or time of the flowering.
23
Except the pollen ovule ratio method, all other methods can lead to erroneous
conclusions, because they are subjective and therefore differ from person to person. For example,
when considering the flower odor, it may be sensed differently by two individuals. Due to this
reason, the results I obtained were found to be different compared to what was found in
literature.
Flower Pollination syndrome Pollination syndrome Literature
according to according to harmonic cited
morphological relation between the pollination
characters flower shape, color and the syndrome
pollinator
Clitoria ternatea Ants Syrphids, bees, butterflies Butterflies
(Katarolu)
24
According to the above comparison, the flowers of Clitoria ternatea employ ants as the
pollination syndrome when determined by using morphological characters, Syrphids, bees and
butterflies according to harmonic relation between the flower shape, color and the pollinator, but
the literature says that Clitoria ternatea flowers are pollinated only by butterflies. Similarly,
other flowers also behave more or less in the same manner.
When we consider the results, some flowers have more than one pollination syndrome,
whereas the others have only one each. The flowers that have more than one pollination
syndrome are called generalized flowers and the flowers having only one pollination syndrome
are called specialized flowers.
The selection of flowers with suitable maturity is extremely important specially when
determining the pollen ovule ratio. Either if the selected flower has already shed pollen or if the
flower is not mature enough the counting of pollens is of less importance. Therefore, flowers
with all pollen retaining inside must be selected.
When determining breeding system by using outcrossing index, to check the stigma
receptivity the stigma was dipped in a solution of hydrogen peroxide (H2O2); if air bubbles are
formed, the stigma was considered to be receptive.
Determining the temporal separation of male and female parts of the flower was also
done under determining breeding system by using outcrossing index, but it requires experience
and skill. According to the results obtained here all the flowers show some potential towards
outbreeding. Outbreeding is the sexual reproduction between individuals whereas inbreeding is
the sexual reproduction within an individual. There are some advantages and disadvantages of
outbreeding and inbreeding:
Advantages of outbreeding
Disadvantages of outbreeding
25
Advantages of inbreeding
Disadvantages of inbreeding
The results obtained from different methods suggest different breeding systems. For
example, according to outcrossing index method Sesbania grandiflora is self compatible and has
some demand for pollinators, but according to pollen ovule ratio method it is an obligate
xenogamous flower. These mismatchings can be due to practical errors.
26
Conlusions
• Tubular, trumpet or bell shaped actinomorphic red, yellow or purple colored and nectar
and nectar guide possessing flowers are pollinated by butterflies and birds.
• Cream or white colored, fragrant, nectar containing nocturnal flowers are pollinated by
moths.
27
References
Holttum, R. E. and I. Enoch. 1991. Gardening in the Tropics. Times Editions Pte Ltd,
Singapore.
http://www.life.uiuc.edu (28/01/2009)
http://www.fs.fed.us (29/01/2009)
http://www.geocities.com (29/01/2009)
http://en.wikipedia.org (29/01/2009)
28