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

On the Genus Euphorbia

This is a guest post. Words and photos by Jeremiah Sandler.

———————

Suspicion

I collect cacti and succulents. The more I collect plants, the more and more I become interested in taxonomic and phylogenetic relationships between them. Not just my own plants – all of them. Most recently, the genus Euphorbia has been on my mind. My favorite species are E. meloformis var. valida and E. horrida.

I’m mostly familiar with the succulent and cacti-looking euphorbia (they are not true cacti) and a few ornamental annuals. Sometimes I would come across a species that I could determine was a euphorbia; but in trying to identify exactly which species, I found countless possibilities within the genus. It seemed odd to me that a single genus could contain so many different forms.

Turns out, Euphorbia consists of over 1800 separate species. What?! That is an insanely high number! Only about 20 genera of plants contain over 1000 separate species. Euphorbia is the fourth most populated genus among all genera of plants.

That staggering number got me thinking: how can a single genus have so many different species? How has the classification worked that out? Has the genus been phylogenetically examined? There’s no way a genus can be so huge. You know what breeders and collectors can do with that much genetic material in a single genus? The man-made hybrids seem endless.

Euphorbia globosa in bloom

Taxonomy

In older taxonomic practices, morphological similarities were the primary method of grouping individuals together. While that is still a common practice today, phylogenetic testing is now an accessible tool for organizing species into related groups.

Organizations such as the Angiosperm Phylogeny Group (APG) have been doing this advanced scientific research – analyzing DNA, doing detailed dissection, etc. Ultimately, they organize plant taxonomy and systematics with greater detail, and examine plant relationships genetically – phylogenetics.

Analyzing genomes is much more expensive and time consuming than observing morphologies. Now, a mix of methods is used, but DNA sequencing has definitely changed the systematics game in a big way. As a result of the APG’s incorporation of widespread phylogenetic DNA analyses, their taxonomical classifications are quickly becoming the generally accepted classifications among plant taxonomists.

Since the inclusion of genetic testing, many plant orders, families, and genera have been reorganized, renamed, expanded, or shrunk.

Euphorbia

One of the identifying features of euphorbias are their very unique flowers. All species in the genus have a cyathium, an inflorescence exclusively produced by euphorbias. Lacking in true petals, sepals, or nectaries, monoecious euphorbia flowers possess only the most essential parts of reproduction. However, bracts, extra-floral nectaries, and other structures surrounding the reproductive parts of the flowers make them appear superficially different.

It would be very time consuming to sequence the DNA of every member of this genus to see where they all fit. Approximately 10% of the euphorbias have been phylogenetically examined, and they confirm the traditional morphological placement. How about that?

Interestingly, of the species genetically analyzed, some were subsequently placed into the genus Euphorbia after historically being considered members of other genera.

Euphorbia horrida and Euphorbia obesa

So? What’s that mean?

Species within the same genus when crossed can (but not always) produce viable offspring. Sometimes they don’t because of differences in pollinators, flowering times, or geographic location, which prevents hybridization. Clades within plant genera also can affect intra-genus reproduction. For example, hard maples won’t naturally hybridize with soft maples, despite both being in the genus Acer. Perhaps the case is similar between the groups within Euphorbia.

As a plant collector and cacti and succulent enthusiast, imagining the endless amounts of hybrids within a massive genus is a fancy idea to me. The APG’s confirming of the initial classifications of Euphorbia into a massive genus makes the idea of endless hybrids all the more real.

Additional guest posts by Jeremiah Sandler:

———————

Jeremiah Sandler lives in southeast Michigan, has a degree in horticultural sciences, and is an ISA certified arborist. Follow him on Instagram: @j.deepsea

Field Trip: Alaska Botanical Garden

While in Anchorage for the Alaska Invasive Species Workshop, I had the chance to visit the Alaska Botanical Garden. As you might expect, the end of October is not the ideal time to be visiting an Alaskan garden, but it was still fun to walk around and imagine what things might look like in their prime while appreciating the year-round beauty that many plants offer.

I arrived on a Saturday morning. The garden was open, but no one else appeared to be around. I walked along the pathways that brought me to all the different cultivated spaces, which cover only a fraction of the 110 acre property. Nervous about bears (signs throughout the garden kept reminding me to be “bear aware”) and wanting to get out of the cold, I skipped the 1.1 mile nature trail that would have taken me around the perimeter of the garden.

While my visit was brief and most of the plants had already gone dormant, I still enjoyed the garden and will make it a point to return if I ever find myself in the area again. In the meantime, here are a few photos I took on that chilly October morning. Apologies in advance as all photos were taken using my cell phone, which is not ideal.

Fruits of highbush cranberry, also known as mooseberry or squashberry (Viburnum edule)

bog rosemary (Andromeda polifolia)

Entrance to the Junior Master Gardener Plot (a.k.a. Children’s Garden)

Ursus botanicus

Astilbe x arendsii ‘Bridal Veil’

alpine cinquefoil (Potentilla villosa)

Entrance to the Herb Garden

Rock Garden maintained by Alaska Rock Garden Society

One of several tufa troughs planted with alpine plants in the Rock Garden

Another tufa trough in the Rock Garden

snowbells (Soldanella sp.)

Saxifraga paniculata var. minutifolia ‘Red-backed Spider’

Holzhaufen or Holz Hausen (a.k.a. German woodpile). Check out this YouTube video to learn how to build your own round woodpile.

———————

More Awkward Botany Field Trips:

Highlights from the Alaska Invasive Species Workshop

This October 24-26th I was in Anchorage, Alaska for the 18th annual Alaska Invasive Species Workshop. The workshop is organized by the Committee for Noxious and Invasive Pests Management and University of Alaska Fairbanks Cooperative Extension. It is a chance for people involved in invasive species management in Alaska – or just interested in the topic – to learn about the latest science, policies, and management efforts within the state and beyond. I am not an Alaska resident – nor had I ever been there until this trip – but my sister lives there, and I was planning a trip to visit her and her family, so why not stop in to see what’s happening with invasive species while I’m at it?

What follows are a few highlights from each of the three days.

Day One

The theme of the workshop was “The Legacy of Biological Invasions.” Ecosystems are shaped by biotic and abiotic events that occurred in the past, both recent and distant. This is their legacy. Events that take place in the present can alter ecosystem legacies. Invasive species, as one speaker said in the introduction, can “break the legacy locks of an ecosystem,” changing population dynamics of native species and altering ecosystem functions for the foreseeable future. Alaska is one of the few places on earth that is relatively pristine, with comparably little human disturbance and few introduced species. Since they are at an early stage in the invasion curve for most things, Alaska is in a unique position to eradicate or contain many invasive species and prevent future introductions. Coming together to address invasive species issues and protect ecosystem legacies will be part of the human legacy in Alaska.

The keynote address was delivered by Jamie Reaser, Executive Director of the National Invasive Species Council and author of several books. She spoke about the Arctic and its vulnerability to invasive species due to increased human activity, climate change, and scant research. To address this and other issues in the Arctic, the Arctic Council put together the Arctic Biodiversity Assessment, and out of that came the Arctic Invasive Alien Species Strategy and Action Plan. Reaser shared some thoughts about how government agencies and conservation groups can come together to share information and how they can work with commercial industries to address the threat of invasive species. She stressed that Alaska can and should play a leadership role in these efforts.

Later, Reaser gave a presentation about the National Invasive Species Council, including its formation and some of the work that it is currently doing. She emphasized that invasive species are a “people issue” – in that the actions and decisions we make both create the problem and address the problem – and by working together “we can do this.”

Day Two

Most of the morning was spent discussing Elodea, Alaska’s first invasive, submerged, freshwater, aquatic plant. While it has likely been in the state for a while, it was only recognized as a problem within the last decade. It is a popular aquarium plant that has been carelessly dumped into lakes and streams. It grows quickly and tolerates very cold temperatures, photosynthesizing under ice and snow. It propagates vegetatively and is spread to new sites by attaching itself to boats and float planes. Its dense growth can crowd out native vegetation and threaten fish habitat, as well as make navigating by boat difficult and landing float planes dangerous. Detailed reports were given about how organizations across the state have been monitoring and managing Elodea populations, including updates on how treatments have worked so far and what is being planned for the future. A bioeconomic risk analysis conducted by Tobias Schwörer was a featured topic of discussion.

After lunch I took a short break from the conference to walk around downtown Anchorage, so I missed a series of talks about environmental DNA. I returned in time to hear an interesting talk about bird vetch (Vicia cracca). Introduced to Alaska as a forage crop, bird vetch has become a problematic weed on farms, orchards, and gardens as well as in natural areas. It is a perennial vine that grows quickly, produces copious seeds, and spreads rhizomatously. Researchers at University of Alaska Fairbanks found that compared to five native legume species, bird vetch produced twice the amount of biomass in the presence of both native and non-native soil microbes, suggesting that bird vetch is superior when it comes to nitrogen fixation. Further investigation found that, using only native nitrogen-fixing bacteria, bird vetch produced significantly more root nodules than a native legume species, indicating that it is highly effective at forming relationships with native soil microbes. Additional studies found that the ability of bird vetch to climb up other plants, thereby gaining access to more sunlight and smothering host plants, contributed to its success as an invasive plant.

 Seed pods of bird vetch (Vicia cracca) in Fairbanks, Alaska

Day Three

The final day of the workshop was a veritable cornucopia of topics, including risk assessments for invasive species, profiles of new invasive species, updates on invasive species control projects, discussions about early detection and rapid response (EDRR), and talks about citizen science and community involvement. My head was swimming with impressions and questions. Clearly there are no easy answers when it comes to invasive species, and like other complex, global issues (made more challenging as more players are involved), the increasingly deep well of issues and concerns to resolve is not likely to ever run dry.

Future posts will dig further into some of the discussions that were had on day three. For now, here are a few resources that I gathered throughout the workshop:

Interpretive sign at Alaska Botanical Garden in Anchorage, Alaska

Drought Tolerant Plants: Water Conservation Landscape at Idaho Botanical Garden

Demonstration gardens are one of the best places to learn about drought tolerant plants that are appropriate for your region. Such gardens not only help you decide which species you should plant, but also show you what the plants look like at maturity, what they are doing at any given time of year, and how to organize them (or how not to organize them, depending on the quality of the garden) in an aesthetically pleasing way. A couple of years ago, I explored the Water Efficient Garden at the Idaho State Capitol Building. This year I visited the Water Conservation Landscape at Idaho Botanical Garden in Boise, Idaho.

The Water Conservation Landscape is planted on a large L-shaped berm on the edge of Idaho Botanical Garden’s property. It is the first thing that visitors to the garden see, before they reach the parking area and the front gate. It is nearly a decade old, so the majority of the plants are well established and in their prime. Because the garden is so visible, year-round interest is important. This imperative has been achieved thanks to thoughtful plant selection and design.

This demonstration garden came about thanks to a partnership between Idaho Botanical Garden and several other organizations, including the water company, sprinkler supply companies, and a landscape designer. An interpretive sign is installed at one end of the garden describing the benefits of using regionally appropriate plants to create beautiful drought tolerant landscapes. If you ever find yourself in the Boise area, this is a garden well worth your visit. In the meantime, here are a few photos as it appeared in 2017.

February 2017

bluebeard (Caryopteris incana ‘Jason’) – February 2017

Sedum spurium ‘Dragon’s Blood – March 2017

winter heath (Erica x darleyensis ‘Kramer’s Red’) – March 2017

May 2017

avens (Geum x hybrida ‘Totally Tangerine’) – May 2017

July 2017

American cranberrybush (Viburnum opulus var. americanum ‘Wentworth’) – July 2017

Fremont’s evening primrose (Oenothera macrocarpa ssp. fremontii ‘Shimmer’) – July 2017

Fremont’s evening primrose (Oenothera macrocarpa ssp. fremontii ‘Shimmer’) – July 2017

August 2017

cheddar pink (Dianthus gratianopolitanus ‘Firewitch’) – August 2017

smoketree (Cotinus coggyria ‘Royal Purple’) – August 2017

gray lavender cotton (Santolina chamaecyparissus) – September 2017

showy stonecrop (Hylotelephium telephium ‘Matrona’) – September 2017

showy stonecrop (Hylotelephium telephium ‘Matrona’) – September 2017

Adam’s needle (Yucca filamentosa ‘Color Guard’) – October 2017

fragrant sumac (Rhus aromatica ‘Gro-Low’) – October 2017

More Drought Tolerant Plant Posts: