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Pollination

Chapter · January 2017

DOI: 10.1016/B978-0-12-809633-8.05070-6

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P.G. Kevan, Pollination in Roses, In Reference Module in Life Sciences, Elsevier, 2017, ISBN: 978-0-12-809633-8,
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Pollination in Roses$
PG Kevan, University of Guelph, Guelph, ON, Canada
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Introduction

Roses attract many insects. Why is this, and what do the insects gain? What do the plants gain? The Rose's Kiss by Peter Bernhardt
provides an entertaining and informative look into the natural history and sex life of flowers, yet information about roses is scarce.
Just what is known about roses and their pollination?
First, it is necessary to define pollination. Pollination is the transfer of pollen from the anthers of a flower to the stigmas of a
different flower on a different plant or on the same plant, or within the same flower. Pollination does not necessarily result in
fertilization of the ovules within the flower's ovary. Self-compatibility is required if self-pollination is to be fruitful.
A pollinator is the agent that causes that transfer. Pollinators range from physical agents, especially the wind (wind pollination
is called anemophily), or biotic agents such as insects, birds, bats and other animals (pollination by insects is called entomophily,
by birds ornithophily, by bats chiropterophily). There is a large technical vocabulary associated with pollination. Roses are
entomophilous. Similarly, plant-breeding systems are highly varied and complex.
When one thinks about insects as visitors to flowers, one thinks of pollinators, nectar and pollen feeders, and sometimes about
feeding damage to flowers. All are part of the anthecology (floral ecology) of roses. The form of the flowers is crucial to
understanding how pollination takes place.

Floral Biology

Flowers of rose species are mostly hermaphroditic and homogamous, meaning that they open with both male (stamens num-
bering from as few as 50 to as many as 250, depending on species) and female (pistils) maturing simultaneously. Rosa setigera
Michx., in which functional male and female flowers are present on different plants (i.e. it is dieocious) is the only exception. In
most species of roses, the flowers are large with rose-colored, white or, less frequently, yellow-colored petals that overlay the much
smaller green sepals that form the sheath of the floral bud. The flowers are referred to as perigynous because the petals and sepals
are borne on a raised, fleshy receptacle, the hip, which surrounds but is not fused with the numerous ovaries that it encloses
(Fig. 1). Most roses have a delightful and characteristic scent. The flowers are devoid of nectar (although some species may secrete
minute amounts on the edge of the receptacle at the base of the flower between the filaments and the ovary) but produce large
amounts of pollen, which is the main reward sought by insect visitors.
That was the state of knowledge when, at the turn of the 20th century, Paul Knuth reviewed what was known about floral
biology in the genus Rosa. He also synthesized the information available on insect visitors to the flowers, noting mostly a wide
range of beetles (Coleoptera), flies (Diptera) and bees (Hymenoptera). As is typical of research in pollination ecology of that day,
little information is given on the role of pollinators in the setting of hips and production of seeds.
Fertilization of the ovules within the hip ensues following pollination; however, the pollen first germinates on the stigma,
grows through the style and liberates its gametes (sperm cells) into the waiting ovule. It may come about through automatic or
insect-mediated selfing within one flower (autogamy), or through pollen transfer from flower to flower on the same plant
(geitenogamy). Cross-pollination (xenogamy) results when pollen is transferred from the anthers of one plant to the stigmas of the
flowers of another.

Self- and Cross-Pollination

One might expect that autogamy in roses, with their mainly homogamous flowers, would be general. Although there are only a
few studies on pollination and breeding systems in roses, it appears that the situation is more complex than originally thought and
that xenogamy is important in several species (Fig. 2). How do the flowers of roses function to favor the transfer of pollen between
flowers and between plants?
The petals of many roses, such as Rosa canina L. and Rosa arvensis Huds., are more or less erect when the flowers first open. The
stamens then spread outwards, exposing the stigmas in the center of the flower. Insect visitors to these flowers tend to land and
perch on the stigmas while feeding at the anthers. As the flowers age over a period of a day, the petals increasingly open, as do the
stamens, and visiting insects land on them and become liberally dusted with pollen. Thus, an insect that has visited an older flower

☆
Change History: February 2016. P.G. Kevan made additions to “Further Reading” section; minor text amendment; correction to first name of photographer of
Fig. 3 (Heidi Dobson).

Reference Module in Life Sciences doi:10.1016/B978-0-12-809633-8.05070-6 1

 Author's personal copy
2 Pollination in Roses

Fig. 1 Rosa rugosa.

Fig. 2 Pollination bag on rose excludes insect- and wind-borne pollen, allows controlled cross- and self-pollinations, and is essential to
understanding the nature of sexual reproductive and floral biology.

readily transfers pollen to a younger one. The mechanism to favor cross-pollination in Rosa rubiginosa L. is further enhanced by the
flowers being slightly (by a matter of an hour or so) protogynous (ie, with the stigmas receptive to pollen germination before the
anthers of the flower have dehisced and started to shed pollen). These mechanisms do not prevent self-pollination, the likelihood
of which would seem to be enhanced in species in which the stamens are tall and curved in over the stigmas, but self-fertilization is
another matter.
The sperm cells in the pollen and egg cells in the ovules are normally produced in plants through a series of cell divisions
involving meiosis and some additional mitotic divisions. As in humans, and most animals and plants, meiotic divisions produce
gametes (sex cells: sperms and eggs) that have half the number of chromosomes (haploid) as the parents (who are diploid). When
a sperm and egg unite during fertilization, the original (diploid) number of chromosomes is restored. During the processes of
meiosis and fertilization, the chromosomes become shuffled and recombined so that variability results. This variability can be
used by plant breeders for making selections, and is used by nature in natural selection for the fittest offspring (ie, those that are
healthiest in turn produce the most and healthiest offspring through Darwinian selection). Self-fertilization severely constrains the
amount of variability of the offspring. The Darwin–Knight law notes that nature disfavors self-pollination and abhors perpetual
self-fertilization.
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Pollination in Roses 3

There are species of roses that are not self-pollinating, some that are weakly self-pollinating, and others that are easily self-
pollinated, but the published data of Eileen Macfarlane are difficult to decipher in regard to the important difference between self-
pollinating and self-fertility, although she wrote: “My experiments showed that the roses are self-fertile.” It is known that two
species she records as highly self-pollinating, R. setigera and Rosa rugosa Thunb., both require cross-pollination by insects to set
fruit. Recently published data indicate that most diploid species, but only a minority of polyploid species, are self-incompatible.
More information is needed about the breeding systems and pollination requirements of the whole genus.
Roses of the section Caninae are polyploids with 2n ¼ 28. In each of these species the pollen contains seven chromosomes
whereas the egg cells contain 2n–7 chromosomes. This situation arises from unusual meiotic divisions which differ on the male
and female side. This strange mechanism may have advantages in providing a way to remove deleterious genes from the genome
(gene complement) that is kept in the seeds. Deleterious genes may be segregated during meiosis to be aborted. Deleterious genes
that are retained during microsporogenesis (pollen production) may be selected against by differential capacity for pollen of
higher genetic quality to be favoured during fertilization. This sort of mate selection is common in plants. It is important for plants
to ensure that seeds in fruits are of the highest quality because investment in the seeds far outweighs that in pollen. Such a strategy
may be especially valuable in maintaining genetic variability and fitness in highly self-fertilizing plants.

Insect Visits, Pollination and Fertilization

Insect visits to roses may be variously important to the plants for sexual reproduction, fruit and seed production, and hence
dispersal. Some roses are self-pollinating and self-fertilizing and can set hips without insect pollination, whereas others cannot.
However, little is known about most species in the genus Rosa. Many of the horticultural varieties do not produce flowers that can
be used by pollinators. Some lack pollen, do not set fruit and must be propagated vegetatively. Some have been so selected for
floral beauty in horticulture that even sexual organs and the characteristic scent are missing.

Floral Advertisements and Rewards for Pollinators

To understand how insects visit the flowers of roses and bring about pollination, one must turn to studies on species, or
wild roses. It is well known that the flowers of wild roses attract a great diversity of insect visitors and almost all feed on pollen.
The wide variety of bees, of which the most conspicuous are bumblebees (Bombus spp.) (Fig. 3), carpenter bees (Xylocopa spp.)
and the familiar honeybees (Apis mellifera), all forage at rose flowers to collect pollen. Bumblebees are often heard making
high-pitched buzzing sounds as they harvest pollen. This sound and vibration serve to release pollen from the anthers. The
bees take the pollen back to their nests where it is used for feeding the larvae. Rose pollen, like pollen of many other flowers, is
highly nutritious, being rich in proteins, amino acids, carbohydrates, lipids and vitamins. Other insects, including some small
bees, eat the pollen while on the flowers. The nutrients in the pollen are converted into yolk in developing eggs of flies, beetles
and other insects. Particularly conspicuous on roses are hover flies (Syrphidae), many of which are beneficial as their larvae
consume aphids.
A wide variety of other insects can be found on the flowers of roses. Most are considered destructive, such as thrips (Thysa-
noptera), chafers (Scarabeidae) and earwigs (Dermaptera). Population studies of thrips on rose flowers allowed the great
Australian ecologists Andrewartha and Birch to model insect population growth over time and under different weather conditions.

Fig. 3 Bombus terrestris visits a flower of Rosa rugosa. Photograph courtesy of Heidi Dobson.
 Author's personal copy
4 Pollination in Roses

Leaf-cutting bees (Megachilidae) sometimes cut circular holes from the petals, but their damage is more commonly seen on the
leaves. These bees are beneficial in nature because of their importance in pollination. The leaf and petal segments they take are
used in the construction of their nests. Crab spiders (Thomisidae) often sit in rose flowers and ambush flower-visiting insects.
The colors of the petals, and the colors of the stamens and pistils, provide a visual target for foraging pollinators. Against the
background of vegetation, the arrangement of colors, mostly pink petals with a center of yellow, is conspicuous to the human eye.
Insects see differently from human beings, having the capacity to see ultraviolet reflections, but mostly not red ones. Insects also have
compound eyes, made of many minute hexagonal facets, that allow them to see shapes in a sort of mosaic. Other differences also
render floral colors difficult to comprehend in the insect world. Green leaves of most plants are uniformly reflecting across the insect
visual spectrum and can be thought of as neutrally colored to pollinators (ie, the equivalent of dull gray). As far as we are aware,
roses do not reflect ultraviolet patterns and the large amount of red reflection is probably not useful to most insects. Thus, most pink
rose petals are usefully reflective to insects in the blue part of the spectrum. The yellow stamens and pistils strongly absorb blue light.
Thus, the petals are colored distinctively from the vegetation, as are the stamens and pistils colored differently from the petals.
Rose flowers tend to be large and it is assumed that they would be visible to pollinators from great distances. This may not be
the case as individual roses are probably invisible to approaching pollinators until they are within about 30–40 cm, perhaps even
much less, as recent research in bee vision physiology suggests.
The odors of roses are produced on various floral parts, sometimes more or less continuously and in some (eg, Rosa damascena
semperflorens) in pulses. The petals are well known for their delightful aroma and are used extensively in the perfume industry. The
best known species for the chemistry of scents in relation to their importance in pollination is R. rugosa. The main chemical is
2-phenyl ethanol with lesser amounts of monoterpenoid alcohols. The anthers and pollen also emit different scents, eugenol and
methyl eugenol, that smell somewhat spicy, and the pollen itself is characteristically scented with long-chain ketones, aldehydes
and esters.
How pollinators orient to and choose flowers at which to forage is complex, involving several sensory systems (vision, scent,
touch and time). Honeybees can distinguish between the male and female flowers of R. setigera, and between newly opened
(brightly colored, pink with yellow anthers) and older flowers (with paler petals and browning anthers). Interchanging fresh petals
on to older flowers did not increase the older flowers' attractiveness, but placing older petals on to fresh flowers did not detract
greatly from floral attractiveness by comparison with that of unmanipulated fresh flowers. Scent differences were not measured.
Bumblebees (Bombus) on R. rugosa (Fig. 3) primarily use the pollen scent to assess the availability of forage in the flowers, but visual
stimuli from the anthers are also important. Petal odors and colors are of secondary use in the bees' selection of rewarding flowers.
Little is known about the role of other floral signals from roses, such as microtextural features that bees can distinguish or
timing of opening. It has been suggested that dish-shaped flowers, such as those of roses, could act as parabolic sonic reflectors that
could amplify the sound of the bees buzzing in the flowers to release pollen and that could be attractive to other bees.

Further Reading

Bernhardt, P., 1999. The Rose's Kiss: A Natural History of Flowers. Washington, DC: Island Press.
Dobson, H.E.M., Bergstrom, G., Groth, I., 1990. Differences in fragrance chemistry between flower parts of Rosa rugosa Thunb (Rosaceae). Israel Journal of Botany 39,
143–156.
Dobson, H.E.M., Danielson, E.M., Van Wesep, I.D., 1999. Pollen odor chemicals as modulators of bumble bee foraging on Rosa rugosa Thunb. (Rosaceae). Plant Species
Biology 14, 153–166.
Faegri, K., van der Pijl, L., 1979. The Principles of Pollination Ecology, third ed. Oxford: Pergamon Press.
Flament, I., Debonneville, C., Furrer, A., 1993. Volatile constituents of roses. In: Teranishi, R., Buttery, R.G., Sugsawa, H. (Eds.), Bioactive Volatile Chemicals from Plants.
American Chemical Society, pp. 269–281. Symposium Series No. 525.
Hosni, K., Zahed, N., Chrif, R., et al., 2011. Volatile oil constituents of Rosa canina L.: Differences related to developmental stages and floral organs. Plant Biosystems 145,
627–634.
Jesse, L.C., Moloney, K.A., Obrycki, J.J, 2006. Insect pollinators of the invasive plant, Rosa multiflora (Rosaceae), in Iowa, USA. Weed Biology and Management 6, 235–240.
Jicinska, D., 1975. Diversity of pollination in some Rosa spp. Preslia (Prague) 47, 267–274.
Jicinska, D., 1976. Autogamy in various species of the genus Rosa. Preslia (Prague) 48, 225–229.
Kemp, J.P., Posluszny, U., Gerrath, J.M., Kevan, P.G., 1993. Floral development of Rosa setigera. Canadian Journal of Botany-Revue Canadienne de Botanique 71, 74–86.
Kemp, J.R., 1994. Floral morphology and pollination biology of Rosa setigera Michaux. PhD Dissertation, University of Guelph, Guelph, ON, Canada.
Kemp, J.R., Kevan, P.G., Posluszny, U., 1993. Morphological differences and changes of the gynoecium in short-lived flowers of Rosa setigera Michaux and their relationship
to dioecy. International Journal of Plant Sciences 154, 550–556.
Knuth, P., 1908. Handbook of Flower Pollination (J.R. Ainsworth Davis, Trans.), vol. II. Oxford: Clarendon Press.
Macfarlane, E.W.E., 1963. A self-pollination mechanism and other items in rose species. American Rose Annual 48, 188–193.
MacPhail, V.J., Kevan, P.G., 2007. Reproductive success and insect visitation in wild roses (Rosa spp.) – Preliminary results from 2004. In: Pemberton, R. (Ed.) Proceedings
of the IVth International Symposium on Rose Research and Cultivation. Acta Horticulturae, vol. 751, pp. 381–388.
Proctor, M., Yeo, P., Lack, A., 1996. The Natural History of Pollination. London: Harper-Collins.
Picone, J.M., Clery, R.A., Watanabe, N., MacTavish, H.S., Turnbull, C.G.N., 2004. Rhythmic emission of floral volatiles from Rosa damascena semperflorens cv. ‘Quatre
Saisons.’ Planta 219, 468–478.
Richards, A.J., 1986. Plant Breeding Systems, second ed. London: George Allen and Unwin.
Ueda, Y., Akimoto, S., 2001. Cross- and self-compatibility in various species of the genus Rosa. Journal of Horticultural Science & Biotechnology 76, 392–395.
Willmer, P., 2011. Pollination and Floral Ecology. Princeton & Oxford: Princeton University Press.
Willson, M.F., 1983. Plant Reproductive Ecology. New York: John Wiley.

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