Daisies: they love me or love me not?

Daisy flowers (Compositae or Asteraceae family) are one of the most complex and evolutionated flowers in the world. In this article, we’ll pick off all the petals from the daisy flowers to understand how this special organ works.

THE ASTERACEAE OR COMPOSITAE: WHO ARE THEY?

The Asteraceae family is the largest family of flowering plants and one of the most worldwide distributed. There are about 25,000 species distributed in 1,100 genera, representing 10% of all plant species currently on earth  and they have a cosmopolitan distribution except the Antarctica.

Many Asteraceae are used on our daily routine. For example there are members of this family in our diet, such as lettuce (Lactuca sativa), chicory or escarole (Cichorium endivia), artichoke (Cynara scolymus) and sunflower (Helianthus annus). Also many species are used in traditional medicine as chamomile (Chamomilla recutita), echinacea (Echinaceae purpurea), dandelion (Taraxacum officinale) or arnica (Arnica montana). They are also many Asteraceae species with horticultural importance, like daisies (Bellis perennis for example, but other species are called so), chrysanthemums (Leucanthemum sp.), marigolds (Calendula sp.) or Dahlias (Dahlia sp.).

Asteraceae species with different uses. A. Dahlia sp., b. Gira-sol (Helianthus annus), c. Arnica montana, d. Echinacea purpurea.

THE CAPITULA

The flower of the Asteraceae species is called capitula and is not a typical flower because it is formed by several flowers grouped together to form a single flower-like structure to attract pollinators. This cluster of flowers imitating a simple flower is called inflorescence. Most Asteraceae present more than one capitula per branch and the way they are organized is structured in a special order. We can found capitulum structured in corimbes or racemes, for example. This structure arranging inflorescences is called a synflorescence.

Normally capitula contain two kinds of flowers: the ray or ligulate flowers and the disc flowers. All have five fused petals.

Ray flower (A), disc flower (B) and schema of the flowers organization in a typical Asteraceae capitula (C), extracted from Greenish (1920).

Ray flowers are usually female flowers, with two connate carpels in an inferior ovary. Their petals are zygomorphic (asymmetrical) and are characterised by the presence of a ligule, a part remembering the typical petal that we pluck off the daisy when playing the game.

Disc flowers are usually hermaphrodites and have a less showy actinomorphic (symmetric) tubular corolla. Disc flowers are in the center of the capitulum looking like small buttons.

The capitula described are the most common in Asteraceae, called heterogamous. The heterogamous capitula can be radiated, as the typical daisy or disciform when only have disc flowers, but the outermost flowers have a long filaments similars to ray flowers, such as in Centaurea sp.

 

Heterogamous disciform capitula of Centaurea deusta in Croacia.

The homogamous capitula have a single type of flower, always hermaphroditic. The discoid homogamous capitula have only disc flowers, like thistles.

Discoid homogamous capitula of Cynara cardunculus.

The ligulate homogamous capitula have only ray flowers, like chicory (Cichorium intybus).

Ligulate homogamous capitula of Cichorium intybus.

CAPITULA ADAPTATIONS

One of the most striking adaptations of the capitulum is that their flowers have different maturation times to avoid self-pollination. The flowers mature centripetally, from the outside in. That’s why we see the disk with o darker color on the inside.

Capitulum of Pericallis echinata, a Canarian endemic, where we can see the diferent degrees of disc flowers maduration.

The secondary pollen presentation is another capitulum adaptation, which is not exclusive from this family but a diagnostic character. The process makes that matured pollen is presented to the pollinators in a different structure from the anthers, the stigma of the pistil, in this case. The secondary pollen presentation occurs by a special adaptation of the anthers, that are fused (syngeneic stamens) forming a tube around the style. Thus, when the mature style is extended through this tube, pollen grains stick making pollen available to pollinators when the stigma reach the outside. This can actually happen because main Asteraceae flowers are proterandrous, i.e. the stamens mature before the style.

Secondary pollen presentation schema (Funk et al., 2009).

This capitulum basic structure has many variations creating many different capitula types. Although most species of Asteraceae are monoecious (we can found hermaphroditic flowers in the same individual) there are dioecious genus, like Baccharis, a genus from tropical South America, which have male and female individuals separately.

Female (left) and male (right) individuals of the dioecious Baccharis sp.

Very rarely, capitula have only a single flower, as in the case of Echinops, where single flowers are grouped in spherical capitula of second order.

Solitary flowers clustered in a second order capitulum of Echinops ritro.

There are another examples of secondary order capitulum, like the famous Edelweiss flower (Leontopodium alpinum). The edelweis flower is particularly interesting because it has densely hairy bracts (with many trichomes) around its discoid capitula acting as white “false” petals and reflecting the high radiation of the high mountains where they live.

Seconsary order capitula of edelweiss (Leontopodium alpinum).

Rarely, capitulum are found alone at the stems apex, not forming synflorescences. This is the case of sunflowers (Helianthus annuus) or Wunderlichia, one of the smallest genus of Asteraceae with six endemic species from Brazil with a really awkward look because of its tomentous indumentum and the lack of leaves when bloom.

Solitary capitulum of Wunderlichia mirabilis in Brazil.

Capitulum pollination is usually made by insects, especially butterflies, which are attracted by the petals color and the nectar, their sweet reward.

The fruit, which is called achene or cypsela in Asteraceae, is formed once the flowers have been fertilized. The cypselae are easy to recognize because many have appendices that look like bristles, awns or scales called pappus acting in wind dispersal.

Diversity of cypselae and pappus found in Asteraceae (Funk et al., 2005).

Now, we can maybe better understand why we can pluck a daisy striping a petals, each from a unique ray flower in the capitulum or why do we blow out so many seeds when we make a wish on a single dandelion flower.

 REFERENCES

• Font Quer P (1953). Diccionario de Botánica. Ed. Labor.
• Funk VA, Bayer RJ, Keeley S, Chan R, Watson L, Emeinholzer B, Schilling E, Panero JL., Baldwin BG, Garcia-Jacas N, Susanna A & Jansen RK (2005). Everywhere but antarctica: using a supertree to understand the diversity and distribution of the Compositae. Biologiske skrifter 55: 343-374.
• Funk VA, Susanna A, Stuessy TF & Bayer RJ (2009). Systematics, evolution, and biogeography of Compositae. International association for plant taxonomy, Vienna, Austria.
• Kadereit JW & Jeffrey C. (2007). The families and genera of vascular plants, vol. 8, Flowering Plants. Eudicots. Asterales. Springer, Berlin.