Types: self-pollination and cross-pollination
An
egg cell in an ovule of a flower may be fertilized by a sperm cell derived from
a pollen grain produced by that same flower or by another flower on the same
plant, in either of which two cases fertilization is said to be due to self-pollination (autogamy);
or, the sperm may be derived from pollen originating on a different plant
individual, in which case the process is called cross-pollination (heterogamy).
Both processes are common, but cross-pollination clearly has certain
evolutionary advantages for the species: the seeds formed may combine the hereditary traits
of both parents, and the resulting offspring generally are more varied than
would be the case after self-pollination. In a changing environment, some of
the individuals resulting from cross-pollination still may be found capable of coping with
their new situation, ensuring survival of the species,
whereas the individuals resulting from self-pollination might all be unable to
adjust. Self-pollination, or selfing, although foolproof in a
stable environment, thus is an evolutionary cul-de-sac. There also is a more
direct, visible difference between selfing and outbreeding (cross-pollination):
in those species where both methods work, cross-pollination usually produces
more, and better quality, seeds. A dramatic demonstration of this effect is
found with hybrid corn (maize), a superior product that results from
cross-breeding of several especially bred lines.
Mechanisms
that prevent self-pollination
STRUCTURAL
Not surprisingly, many species of plants have developed mechanisms
that prevent self-pollination. Some—e.g., date palms (Phoenix
dactylifera) and willows (Salix species)—have become dioecious;
that is, some plants produce only “male” (staminate) flowers, with the rest
producing only “female” (pistillate or ovule-producing) ones. In species in
which staminate and pistillate flowers are found on the same individual (monoecious plants)
and in those with hermaphroditic flowers (flowers possessing both stamens and
pistils), a common way of preventing self-fertilization is to have the pollen
shed either before or after the period during which the stigmas on the same
plant are receptive, a situation known as dichogamy. The more usual form of dichogamy, which is found
especially in such insect-pollinated flowers as fireweed (Epilobium
angustifolium) and salvias (Salvia species), is protandry, in which the stamens ripen before the
pistils. Protogyny,
the situation in which the pistils mature first, occurs in arum lilies and many
wind-pollinated plants, such as grasses—although several grasses are
self-pollinated, including common varieties of wheat,
barley, and oats. Avocado has both protogynous and protandrous varieties, and
these often are grown together to encourage cross-fertilization. A structural feature of flowers
that discourages selfing is heterostyly, or variation in the length of thestyle (neck
of the pistil). This occurs in the common primrose (Primula
vulgaris) and species of woodsorrel (Oxalis) and flax.
In most British primrose populations, for example, approximately half the
individuals have so-called “pin” flowers, which possess
short stamens and a long style, giving the stigma a position at the flower’s
mouth, whereas the other half have “thrum” flowers, in
which the style is short and the stamens are long, forming a “thrumhead” at the opening of the flower. Bees can hardly
fail to deposit the pollen they receive from one type of flower onto the
stigmas of the other type. The genetic system that regulates flower structure
in these primroses is so constituted that cross-pollination automatically
maintains a 50:50 ratio between pins and thrums. In the flowers of purple loosestrife(Lythrum salicaria),
the stamens and styles are of three different lengths to limit self-fertilization.
CHEMICAL
Chemical self-incompatibility is
another device for preventing self-fertilization. In this phenomenon, which
depends on chemical substances within the plant, the pollen may fail to grow on
a stigma of the same flower that produced it or, after germination, the pollen
tube may not grow normally down the style to effect fertilization. The process
is controlled genetically; it need not be absolute and can change in degree
during the flowering season. Not surprisingly, chemical incompatibility usually
is not found in those plants that have strong structural or temporal barriers
against self-pollination. Formation of one such mechanism during evolution
apparently was enough for most plant species.
Mechanisms
that permit self-pollination
In many instances, successful self-pollination takes place at the end
of a flower’s life-span if cross-pollination has not occurred. Such
self-pollination may be achieved by curving of stamens or style as occurs, for
example, in fireweed. It can be an evolutionary advantage when animal pollinators are temporarily scarce or
when the plants in a population are widely scattered. Under such circumstances,
selfing may tide the species over until better circumstances for outbreeding
arrive. For this reason, selfing is common among annual plants; these often must produce an
abundance of seed for the rapid and massive colonization of any bare ground
that becomes available. If, in a given year, an annual plant were to produce no
seed at all, survival of the species might be endangered. A persistent habit of
self-pollination apparently has been adopted successfully by some plant species
whose natural pollinators have died out. Continued selfing also is practiced by
many food-crop plants. Some of these plants are cleistogamous,
meaning that the flowers fail to open, an extreme way of ensuring
self-pollination. A similar process is apomixis, the development of an ovule into a seed
without fertilization. Apomixis is easily demonstrated in lawn dandelions,
which produce seeds even when stamens and styles are cut off just before the
flowers open. Consistent apomixis has the same pros and cons as continued
selfing. The offspring show very little genetic variability, but there is good
survival if the species is well adapted to its habitat and if the environment
does not change.
Agents of
pollen dispersal
Insects
BEETLES AND FLIES
The ancient principle of trapping insects as a means of ensuring
pollination was readopted by some advanced families (e.g., orchids and
milkweeds), and further elaboration perfected the flower traps of
primitive families. The cuckoopint (Arum
maculatum), for example, attracts minute flies, which normally
breed in cow dung, by means of a fetid smell. This smell is generated in early
evening, along with considerable heat, which helps to volatilize the odour
ingredients. The flies visiting the plant, many of which carry Arum pollen, enter the floral trap through
a zone of bristles and then fall into a smooth-walled floral chamber from which
escape is impossible. Gorging themselves on a nutritious stigmatic secretion
produced by the female flowers at the base of the chamber, the flies effect
cross-pollination. Late at night, when the stigmas no longer function, the male
flowers, situated much higher on the floral column, proceed to bombard the
flies with a rain of pollen. The next day, when smell, heat, and food are gone,
the prisoners, “tarred” with stigmatic secretion and “feathered” with pollen,
are allowed to escape by a wilting of the inflorescence (flower
cluster). Usually the escaped flies are soon recaught by another inflorescence,
which is still in the smelly, receptive stage, and cross-pollination again
ensues. Superb timing mechanisms underlie these events. The heat-generating
metabolic process in the inflorescence is triggered by a hormone, calorigen, originating
in the male flower buds only under the right conditions. The giant
inflorescences of the tropical plant Amorphophallus
titanumsimilarly trap large carrion beetles.
In general, trap flowers victimize beetles or flies of a primitive
type. Although beetles most likely were involved as pollinators when flowering
plants as a group were born, their later performance in pollination has been
disappointing. Some modern beetles do visit smelly flowers of an open type,
such as elderberry and hawthorn, but with few exceptions they are still mainly
pollen eaters. Flies as a group have become much more diversified in their
habits than beetles have. Female short-tongued flies may be deceived by
open-type flowers with carrion smells—e.g., the flowers of Stapelia and Rafflesia.Mosquitoes with
their long tongues are effective pollinators of certain orchids (Habenaria species) in North American swamps. In
Europe, the bee fly (Bombylius) is an important long-tongued
pollinator. Extremely specialized as nectar drinkers are certain South African
flies; for example, Megistorhynchus longirostris,
which has a tongue that is 60 to 70 mm (2.3 to 2.7 inches) long.
The
voraciousness of flower beetles demonstrates the futility of enticing insect
pollinators solely with such an indispensable material as pollen. As a
defensive strategy, certain nectar-free flowers that cater to beetles and
bees—such as wild roses, peonies, and poppies—produce a superabundance of
pollen. Other plants—e.g., Cassia—have
two types of stamens, one producing a special sterile pollen used by insects as
food, the other yielding normal pollen for fertilizing the ovules. Other
flowers contain hairs or food bodies that are attractive to insects.
BEES
In the modern world, bees are probably the most important insect
pollinators. Living almost exclusively on nectar, they feed their larvae pollen
and honey (a modified nectar). To obtain their foods, they possess striking
physical and behavioral adaptations, such as tongues as long as 2.5 cm (1
inch), hairy bodies, and (in honeybees and
bumblebees) special pollen baskets. The Austrian naturalist Karl von Frisch has
demonstrated that honeybees, although blind to red light, distinguish at least
four different colour regions, namely, yellow (including
orange and yellow green), blue green, blue (including purple and violet), and
ultraviolet. Their sensitivity to ultraviolet enables bees to follow
nectar-guide patterns not apparent to the human eye. They are able to taste
several different sugars and also can be trained to differentiate between
aromatic, sweet, or minty odours but not foul smells. Fragrance may be the
decisive factor in establishing the honeybee’s habit of staying with one
species of flower as long as it is abundantly available. Also important is that honeybee workers can communicate to one another
both the distance and the direction of an abundant food source by means of
special dances.
Bee flowers, open in the daytime, attract their insect
visitors primarily by bright colours; at close range, special patterns and
fragrances come into play. Many bee flowers provide their visitors with a
landing platform in the form of a broad lower lip on which the bee sits down
before pushing its way into the flower’s interior, which usually contains both
stamens and pistils. The hermaphroditism of most bee flowers makes for
efficiency, because the flower both delivers and receives a load of pollen
during a single visit of the pollinator, and the pollinator never travels from
one flower to another without a full load of pollen. Indeed, the floral
mechanism of many bee flowers permits only one pollination visit. The pollen
grains of most bee flowers are sticky, spiny, or highly sculptured, ensuring
their adherence to the bodies of the bees. Since one load of pollen contains
enough pollen grains to initiate fertilization of many ovules, most individual
bee flowers produce many seeds.
Examples of flowers that depend heavily on bees are larkspur,
monkshood, bleeding heart, and Scotch broom. Alkali bees (Nomia) andleaf-cutter bees (Megachile)
are both efficient pollinators of alfalfa; unlike honeybees, they are not
afraid to trigger the explosive mechanism that liberates a cloud of pollen in alfalfa flowers. Certain Ecuadorian orchids (Oncidium) are
pollinated by male bees of the genus Centris; vibrating in
the breeze, the beelike flowers are attacked headlong by the strongly
territorial males, who mistake them for competitors. Other South American
orchids, nectarless but very fragrant, are visited by male bees (Euglossa species) who, for reasons not yet
understood, collect from the surface of the flowers an odour substance, which
they store in the inflated parts of their hindlegs.
WASPS
Few wasps
feed their young pollen or nectar. Yellow jackets, however, occurring occasionally in
large numbers and visiting flowers for nectar for their own consumption, may
assume local importance as pollinators. These insects prefer brownish-purple flowers with
easily accessible nectar, such as those of figwort. The flowers of some
Mediterranean and Australian orchids mimic the females of certain wasps (of the
families Scoliidae and Ichneumonidae) so successfully that the males of these species
attempt copulation and receive the pollen masses on their bodies. In figs, it
is not the pollinator’s sexual drive that is harnessed by the plant but the
instinct to take care of the young; tiny gall wasps (Blastophaga)
use the diminutive flowers (within their fleshy receptacles) as incubators.
BUTTERFLIES AND MOTHS
The evolution of moths and butterflies (Lepidoptera) was made
possible only by the development of the modern flower, which provides their
food. Nearly all species of Lepidoptera have a tongue, or proboscis, especially
adapted for sucking. The proboscis is coiled at rest and extended in feeding. Hawkmoths hover while they feed, whereas
butterflies alight on the flower. Significantly, some butterflies can taste
sugar solutions with their feet. Although moths, in general, are nocturnal and
butterflies are diurnal, a colour sense has been demonstrated in representatives
of both. Generally, the colour sense in Lepidoptera is similar to that of bees,
but swallowtails and certain other butterflies also respond to red colours.
Typically, colour and fragrance cooperate in guiding Lepidoptera to flowers,
but in some cases there is a strong emphasis on just one attractant; for
example, certain hawkmoths can find fragrant honeysuckles hidden from sight.
Typical moth
flowers—e.g., jimsonweed, stephanotis, and honeysuckle—are light-coloured,
often long and narrow, without landing platforms. The petals are sometimes
fringed; the copious nectar is often in a spur. They are open and
overwhelmingly fragrant at night. Butterfly flowers—e.g., those of butterfly bush,
milkweed, and verbena—are conspicuously coloured, often red, generally smaller
than moth flowers, but grouped together in erect, flat-topped inflorescences
that provide landing space for the butterflies.
Important
pollinating moths are the various species of the genus Plusia, sometimes
occurring in enormous numbers, and the hummingbird hawkmoth (Macroglossa), which
is active in daylight. A small moth, Tegeticula maculata, presents an
interesting case. It is totally dependent on yucca flowers, in whose ovules its
larvae develop. Before depositing their eggs, the females pollinate the
flowers, following an almost unbelievable pattern of specialized behaviour,
which includes preparing a ball of pollen grains and carrying it to the stigma
of the plant they are about to use for egg laying.
Wind
Although
prevalent in the primitive cycads and in conifers, such as pine and fir, wind
pollination (anemophily) in the flowering plants must be considered as a
secondary development. It most likely arose when such plants left the tropical
rain forest where they originated and faced a more
hostile environment, in which the wind weakened the effectiveness of smell as
an insect attractant and the lack of pollinating flies and beetles also made
itself felt. Lacking in precision, wind pollination is a wasteful process. For
example, one male plant of Mercurialis
annua, a common weed, produces 1.25 billion grains of pollen to be
dispersed by the wind; a male sorrel plant produces 400 million. Although, in
general, the concentration of such pollen becomes very low about one-fourth
mile (0.4 km) from its source, nonetheless in windy areas it can cover
considerable distances. Pine pollen, for example, which is naturally equipped
with air sacs, can travel up to 500 miles (800 km) although the grains may lose
their viability in the process. Statistically, this still gives only a slim
chance that an individual stigma will be hit by more than one or two pollen
grains. Also relevant to the number of pollen grains per stigma is the fact
that the dry, glueless, and smooth-surfaced grains are shed singly. Since the number
of fertilizing pollen grains is low, the number of ovules in a single flower is
low and, as a consequence, so is the number of seeds in each fruit. In hazel,
walnut, beech, and oak, for example, there are only two ovules per flower, and,
in stinging nettle, elm, birch, sweet gale, and grasses, there is only one. Wind-pollinated
flowers are inconspicuous, being devoid of insect attractants and rewards, such
as fragrance, showy petals, and nectar. To facilitate exposure of the flowers
to the wind, blooming often takes place before the leaves are out in spring, or
the flowers may be placed very high on the plant. Inflorescences, flowers, or
the stamens themselves move easily in the breeze, shaking out the pollen, or
the pollen containers (anthers)
burst open in an explosive fashion when the sun hits them, scattering the
pollen widely into the air. The stigmas often are long and divided into arms or
lobes, so that a large area is available for catching pollen grains. Moreover,
in open areas wind-pollinated plants of one species often grow together in
dense populations. The chance of self-pollination, high by the very nature of
wind pollination, is minimized by the fact that many species are dioecious or (like
hazel) have separate male and female flowers on each plant. Familiar flowering
plants relying on wind pollination are grasses, rushes, sedges, cattail,
sorrel, lamb’s-quarters, hemp, nettle, plantain, alder, hazel, birch, poplar,
and oak. (Tropical oaks, however, may be insect-pollinated.)
Birds
Because the study of mechanisms of pollination began in Europe, where
pollinating birds are rare, their importance is often underestimated. In fact,
in the tropics and the southern temperate zones, birds are at least as
important as pollinators as insects are, perhaps more so. About a third of the
300 families of flowering plants have at least some members with ornithophilous
(“bird-loving”) flowers—i.e., flowers attractive to birds. Conversely, about
2,000 species of birds, belonging to 50 or more families, visit flowers more or
less regularly to feed on nectar, pollen, and flower-inhabiting insects or
spiders. Special adaptations to this way of life, in the form of slender,
sometimes curved, beaks and tongues provided with brushes or shaped into tubes,
are found in over 1,600 species of eight families:hummingbirds, sunbirds (see The Rodent That Acts Like a Hippo and Other Examples of Convergent
Evolution), honeyeaters,
brush-tongued parrots, white-eyes, flower-peckers, honeycreepers (or sugarbirds), and Hawaiian honeycreepers such as the iiwi.
Generally, the sense of smell in birds is
poorly developed and not used in their quest for food; instead, they rely on
their powerful vision and their colour sense, which resembles that of human
(ultraviolet not being seen as a colour, whereas red is). Furthermore, the
sensitivity of the bird’s eye is
greatest in the middle and red part of the spectrum. This is sometimes ascribed
to the presence in the retina of orange-red drops of oil, which together may
act as a light filter.
Although other explanations have been forwarded,
the special red sensitivity of the bird eye is usually thought to be the reason
why so many bird-pollinated flowers are of a uniform, pure red colour.
Combinations of complementary colors, such as orange and blue, or green and
red, also are found, as are white flowers. As might be expected, bird flowers
generally lack smell and are open in the daytime; they are bigger than most
insect flowers and have a wider floral tube. Bird flowers also are sturdily
constructed as a protection against the probing bill of the visitors, with the
ovules kept out of harm’s way in an inferior ovary beneath the floral chamber
or placed at the end of a special stalk or behind a screen formed by the fused
bases of the stamens. The latter, often so strong as to resemble metal wire,
are usually numerous, brightly coloured, and protruding, so that they touch a
visiting bird on the breast or head as it feeds. The pollen grains often stick
together in clumps or chains, with the result that a single visit may result in
the fertilization of hundreds of ovules.
In the
Americas, where hummingbirds usually
suck the nectar of flowers on the wing, ornithophilous flowers (e.g., fuchsias)
are often pendant and radially symmetrical, lacking the landing platform of the
typical bee flower. In Africa and Asia, bird flowers often are erect and do
offer their visitors, which do not hover, either a landing platform or special
perches in the form of small twigs near the flower . Pollinating birds are
bigger than insects and have a very high rate of metabolism. Although some
hummingbirds go into a state resembling hibernation every night, curtailing
their metabolism drastically, others keep late hours. Thus, in general, birds
need much more nectar per individual than insects do. Accordingly, bird flowers
produce nectar copiously—a thimbleful in each flower of the coral tree, for
example, and as much as a liqueur glassful in flowers of the spear lily (Doryanthus).
Plants bearing typical bird flowers are cardinal flower, fuchsia, red
columbine, trumpet vine, hibiscus, strelitzia, and eucalyptus, and many members
of the pea, orchid, cactus, and pineapple families.
Mammals
In Madagascar,
the mouse lemurs (Microcebus), which are only ten
centimetres (four inches) long, obtain food from flowers, and in Australia the
diminutive marsupial honey possums and pygmy possums also are flower
specialists. Certain highly specialized tropical bats,
particularlyMacroglossus and Glossophaga,
also obtain most or all of their food from flowers. The Macroglossus (big-tongued) species of southern Asia
and the Pacific are small bats with sharp snouts and long, extensible tongues,
which carry special projections (papillae) and sometimes a brushlike tip for
picking up a sticky mixture of nectar and pollen. Significantly, they are
almost toothless. Colour sense and that sonar sense so prominent in other bats,
seem to be lacking. Their eyesight is keen but, since they feed at night, they
are probably guided to the flowers principally by their highly developed sense
of smell. The bats hook themselves into the petals with their thumb claws and
stick their slender heads into the flowers, extracting viscid nectar and
protein-rich pollen with their tongues. The plants involved have, in the
process of evolution, responded to the bats by producing large
(sometimes huge) amounts of these foods. One balsa-tree flower, for example,
may contain a full 10 grams (0.3 ounce) of nectar, and one flower from a baobab
tree has about 2,000 pollen-producing stamens. Some bat flowers also provide succulent petals
or special food bodies to their visitors. Another striking adaptation is that
the flowers are often placed on the main trunk or the big limbs of a tree (cauliflory); or, borne on thin, ropelike branches, they
dangle beneath the crown (flagelliflory). The pagoda
shape of the kapok tree serves the same purpose: facilitation of the bat’s
approach. Characteristics of the flowers themselves include drab colour, large
size, sturdiness, bell-shape with wide mouth and, frequently, a powerful rancid
or urinelike smell.
The giant saguaro cactus and the century plant (Agave) are
pollinated by bats, although not exclusively, and cup-and-saucer vine (Cobaea
scandens) is the direct descendant of a bat-pollinated American
plant. Calabash, candle tree, and areca palm also have bat-pollinated flowers.
Water
Although
pollen grains can be made to germinate in aqueous sugar solutions, water alone
in most cases has a disastrous effect on them. Accordingly, only a very few
terrestrial plants, such as the bog asphodel of the Faroes, use rainwater as a
means of pollen transport. Even in aquatic plants, water is seldom the true
medium of pollen dispersal. Thus, the famous Podostemonaceae, plants that grow only on rocks in
rushing water, flower in the dry season when the plants are exposed;
pollination occurs with the aid of wind or insects or by selfing. Another
aquatic plant, ribbon
weed, sends its male and female flowers to the surface separately. There, the
former transform themselves into minute sailboats, which are driven by the wind
until they collide with the female flowers. In the Canadian waterweed, and also
in pondweed (Potamogeton) and ditch grass (Ruppia),
the pollen itself is dispersed on the water’s surface; it is, however, still
water-repellent. True water dispersal (hydrophily), in which the pollen grains
are wet by water, is found only in the hornworts and eelgrasses.