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.

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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:

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

Maize Anatomy and the Anatomy of a Maze

Commonly known as corn throughout much of North America, maize is a distinctive emblem of the harvest season. It is one of the most economically important crops in the world (the third most important cereal after rice and wheat) and has scads of uses from food to feed to fuel. The story of its domestication serves as a symbol of human ingenuity, and its plasticity in both form and utility is a remarkable example of why plants are so incredible.

The genus Zea is in the grass family (Poaceae) and consists of five species: Z. diploperennis, Z. perennis, Z. luxurians, Z. nicaraguensis, and Z. mays. Maize is the common name of Zea mays subsp. mays, which is one of four Z. mays subspecies and the only domesticated taxon in the genus. All other taxa are commonly and collectively referred to as teosintes.

The domestication of maize, apart from being an impressive feat, has long been a topic of research and a challenging story to tease apart. The current understanding is that maize was first domesticated around 9000 years ago in the Balsas River valley in southern Mexico, the main progenitor being Zea mays subsp. parviglumis. It is astonishing how drastically different in appearance teosintes are from modern day maize, but it also explains why determining the crop wild relative of maize was so difficult.

Teosinte, teosinte-maize hybrid, and maize - photo credit: wikimedia commons

Teosinte, teosinte-maize hybrid, and maize – photo credit: wikimedia commons

Teosintes and maize both have tall central stalks; however, teosintes generally have multiple lateral branches which give them a more shrubby appearance. In teosinte, each of the lateral branches and the central stalk terminate in a cluster of male flowers; female flowers are produced at the nodes along the lateral branches. In maize, male flowers are borne at the top of the central stalk, and lateral branches are replaced by short stems that terminate in female flowers. This is where the ears develop.

Ears – or clusters of fruits – are blatantly different between teosintes and maize. To start with, teosinte produces a mere 5 to 12 fruits along a short, narrow cob (flower stalk). The fruits are angular and surrounded in a hard casing. Maize cobs are considerably larger both in length and girth and are covered in as many as 500 or more fruits (or kernels), which are generally more rounded and have a softer casing. They also remain on the cob when they are ripe, compared to teosinte ears, which shatter.

Evolutionary biologist, Sean B. Carroll, writes in a New York Times article about the amazing task of “transform[ing] a grass with many inconvenient, unwanted features into a high-yielding, easily harvested food crop.” These “early cultivators had to notice among their stands of plants variants in which the nutritious kernels were at least partially exposed, or whose ears held together better, or that had more rows of kernels, and they had to selectively breed them.” Carroll explains that this “initial domestication process which produced the basic maize form” would have taken several hundred to a few thousand years. The maize that we know and love today is a much different plant than its ancestors, and it is still undergoing regular selection for traits that we find desirable.

Female inflorescence (or "ear") of Zea mays subsp. mays - photo credit: wikimedia commons

Female inflorescence (or “ear”) of Zea mays subsp. mays – photo credit: wikimedia commons

To better understand and appreciate this process, it helps to have a basic grasp of maize anatomy. Maize is an impressive grass in that it regularly reaches from 6 to 10 feet tall and sometimes much taller. It is shallow rooted, but is held up by prop or brace roots – adventitious roots that emerge near the base of the main stalk. The stalk is divided into sections called internodes, and at each node a leaf forms. Leaf sheaths wrap around the entirety of the stalk, and leaf blades are long, broad, and alternately arranged. Each leaf has a prominent midrib. The stalk terminates in a many-branched inflorescence called a tassel.

Maize Anatomy 101 - image credit: Canadian Goverment

Maize Anatomy 101 – image credit: Canadian Government

Maize is monoecious, which means that it has separate male and female flowers that occur on the same plant. The tassel is where the male flowers are located. A series of spikelets occur along both the central branch and the lateral branches of the tassel. A spikelet consists of a pair of bracts called glumes, upper and lower lemmas and paleas (which are also bracts), and two simple florets composed of prominent stamens. The tassel produces and sheds tens of thousands of pollen grains which are dispersed by wind and gravity to the female inflorescences below and to neighboring plants.

Female inflorescences (ears) occur at the top of short stems that originate from leaf axils in the midsection of the stalk. Leaves that develop along this reduced stem wrap around the ears forming the husk. Spikelets form in rows along the flower stalk (cob) within the husk. The florets of these spikelets produce long styles that extend beyond the top of the husk. This cluster of styles is known as the silk. When pollen grains land on silk stigmas, pollen tubes grow down the entire length of the silks to reach the embryo sac. Successful fertilization produces a kernel.

The kernel – or fruit – is known botanically as a caryopsis, which is the standard fruit type of the grass family. Because the fruit wall and seed are fused together so tightly, maize kernels are commonly referred to as seeds. The entire plant can be used to produce feed for animals, but it is the kernel that is generally consumed (in innumerable ways) by humans.

There is so much more to be said about maize. It’s a lot to take in. Rather than delve too much further at this point, let’s explore one of the other ways that maize is used by humans to create something that has become another feature of the fall season – the corn maze.

Entering the corn maze at The Farmstead in Meridian, Idaho

Exploring the corn maze at The Farmstead in Meridian, Idaho

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