The Agents That Shape the Floral Traits of Sunflowers

Flowers come in a wide array of shapes, sizes, colors, and scents. Their diversity is downright astounding. Each individual species of flowering plant has its own lengthy story to tell detailing how it came to look and act the way it does. This is its evolutionary history. Unraveling this history is a nearly insurmountable task, but one that scientists continue to chip away at piece by piece.

In the case of floral traits – particularly for flowers that rely on pollinators to produce seeds – it is safe to say that millennia of interactions with floral visitors have helped shape not only the way the flower looks, but also the nature of its nectar and pollen. However, flowers are “expensive” to make and maintain, so even though they are necessary for reproduction, plants must find a balance between that and allocating resources for defense – against both herbivory and disease – and growth. This balance can differ depending on a plant’s life history – whether it is annual or perennial. An annual plant has one shot at reproduction, so it can afford to funnel much of its energy there. If a perennial is unsuccessful at reproduction one year, there is always next year, as long as it has allocated sufficient resources towards staying alive.

Where a plant exists in the world also influences how it looks. Abiotic factors like temperature, soil type, nutrient availability, sun exposure, and precipitation patterns help shape, through natural selection, many aspects of a plant’s anatomy and physiology, including the structure and composition of its flowers. Additional biotic agents like nectar robbersflorivores, and pathogens can also influence certain floral traits.

This is the background that researchers from the University of Central Florida and University of Georgia drew from when they set out to investigate the reasons for the diverse floral morphologies in the genus Helianthus. Commonly known as sunflowers, Helianthus is a familiar genus consisting of more than 50 species, most of which are found across North America. The genus includes both annuals and perennials, and all but one species rely on cross-pollination to produce viable seeds. Pollination is mainly carried out by generalist bees.

Maximilian sunflower (Helianthus maximiliani)

Helianthus species are found in diverse habitats, including deserts, wetlands, prairies, rock outcrops, and sand dunes. Their inflorescences – characteristic of plants in the family Asteraceae – consist of a collection of small disc florets surrounded by a series of ray florets, which as a unit are casually referred to as a single flower. In Helianthus, ray florets are completely sterile and serve only to attract pollinators. Producing large and numerous ray florets takes resources away from the production of fertile disc florets, and sunflower species vary in the amount of resources they allocate for each floret form.

In a paper published in the July 2017 issue of Plant Ecology and Evolution, researchers selected 27 Helianthus species and one Phoebanthus species (a closely related genus) to investigate “the evolution of floral trait variation” by examining “the role of environmental variation, plant life history, and flowering phenology.” Seeds from multiple populations of each species were obtained, with populations being carefully selected so that there would be representations of each species from across their geographic ranges. The seeds were then grown out in a controlled environment, and a series of morphological and physiological data were recorded for the flowers of each plant. Climate data and soil characteristics were obtained for each of the population sites, and flowering period for each species was collected from various sources.

The researchers found “all floral traits” of the sunflower species to be “highly evolutionarily labile.” Flower size was found to be larger in regions with greater soil fertility, consistent with the resource-cost hypothesis which “predicts that larger and more conspicuous flowers should be selected against in resource-poor environments.” However, larger flower size had also repeatedly evolved in drier environments, which goes against this prediction. Apart from producing smaller flowers in dry habitats, flowering plants have other strategies to conserve water such as opening their flowers at night or flowering for a short period of time. Sunflowers do neither of these things. As the researchers state, “this inconsistency warrants consideration.”

The researchers speculate that “the evolution of larger flowers in drier environments” may be a result of fewer pollinators in these habitats “strongly favoring larger display sizes in self-incompatible species.” The flowers are big because they have to attract a limited number of pollinating insects. Conversely, flowers may be smaller in wetter environments because there is greater risk of pests and diseases. This is supported by the enemy-escape hypothesis – smaller flowers are predicted in places where there is increased potential for florivory and pathogens. Researchers found that lower disc water content had also evolved in wetter environments, which supports the idea that the plants may be defending themselves against flower-eating pests.

Seed heads of Maximilian sunflower (Helianthus maximiliani)

Another interesting finding is that, unlike other genera, annual and perennial sunflower species allocate a similar amount of resources towards reproduction. On average, flower size was not found to be different between annual and perennial species. Perhaps annuals instead produce more flowers compared to perennials, or maybe they flower for longer periods. This is something the researchers did not investigate.

Finally, abiotic factors were not found to have any influence on the relative investment of ray to disc florets or the color of disc florets. Variations in these traits may be influenced instead by pollinators, the “biotic factor” that is considered “the classic driver of floral evolution.” This is something that will require further investigation. As the researchers conclude, “determining the exact drivers of floral trait evolution is a complex endeavor;” however, their study found “reasonable support for the role of aridity and soil fertility in the evolution of floral size and water content.” Yet another important piece to the puzzle as we learn to tell the evolutionary history of sunflowers.

Advertisements

Summer of Weeds: Flower of an Hour

Hibiscus trionum is a great example of an ornamental plant becoming a widespread weed. Its common name, flower of an hour, refers to its short-lived blooms. Other common names include Venice mallow, bladder hibiscus, bladderweed, modesty, and shoofly. Native to southern Europe and tropical to subtropical parts of Asia and Africa, it was introduced to America as an attractive addition to annual flower beds. It is now naturalized in many states across the country.

Hibiscus is a huge genus in the family Malvaceae, consisting of species found throughout warmer parts of the world. H. trionum is a warm season annual that grows to around two feet tall and has the habit of a sprawling, decumbent vine; an upright, many-branched mound; or something in-between. Its leaves are alternately arranged and three-lobed with coarsely toothed margins. The flowers are solitary and borne in the axils of leaves. They are creamy white to pale yellow with a purple-brown center, and are both cross- and self-pollinated.

flower of an hour (Hibiscus trionum)

Flowering occurs on sunny days throughout the summer. The ephemeral flowers promptly produce a balloon-shaped seed capsule that is hairy and papery with prominent purple veins. Once mature, the capsules split open at the top to reveal five compartments lined with brown to black, kidney- or heart-shaped seeds. Every part of this plant is attractive and interesting to look at, which is why it is no surprise that it is welcome in many flower beds.

Seed capsule of flower of an hour (Hibiscus trionum)

Sites that are in full sun with fertile soil and regular moisture are sought after by flower of an hour. Less fertile soils are still prone to invasion. As with many weeds, disturbance is key, so it is often found in agricultural fields, rangelands, along roadsides, and in vacant lots and construction sites. Its presence in natural areas is a result of escaping from garden beds, agricultural fields, etc.

When we choose to grow plants that have a history of escaping into natural areas, we should be aware of both our proximity to natural areas and the dispersal mechanisms of the plants. Exotic plants that reproduce reliably and prolifically by seed, such as flower of an hour, should be considered unsuitable for gardens that are adjacent to natural areas.

This is because many popular ornamental plants have become invasive in the wild. Plants that are perfectly welcome in our gardens manage to find suitable habitat in natural areas, potentially threatening the livelihood of native plants and/or altering ecological processes such as fire regimes. An example of this where I live is bachelor’s buttons (Centaurea cyanus), which has escaped from gardens and invaded the Boise Foothills. While the impact of this invasion is not well-studied, the speed at which this plant has spread is disconcerting. Even more disconcerting is the fact that seeds of this and other European and Asian species are commonly found in “wildflower” seed mixes distributed throughout North America.

While I am sympathetic towards weeds, I also see them as one of the best reminders of the impacts that humans can have on the planet. They are clear indicators that every step we take has consequences. We should be mindful of this, and we should continue to have the tough conversations that issues like weeds and their impacts encourage us to have. There are no easy answers, but the dialogue must go on. Because all of us – gardeners and non-gardeners/ecologists and non-ecologists alike – generally have an opinion about weeds, they seem like a pretty good place to start.

Additional Resources:

Quote of the Week:

From the book Invasive Plant Medicine by Timothy Lee Scott

The nature of a weed is opportunistic, and we, as humans, have created enormous holes of opportunity for these plants to fill. They have adapted to be at our side, waiting for those favorable times to cover the exposed soils that we continually create. With ever-changing genetics of form, function, and transmutation, weeds have evolved to withstand the punishments that humans unleash upon them.