The Wonderful World of Plantlets, Bulbils, Cormlets, Tubercles, and Gemmae

Probably the most well known strategy that plants have for dispersal is by way of seeds. Seeds are plants in embryo, and new generations of plants are born when seeds, released from their parent plants, find suitable locations to germinate. But one of the most amazing things about plants in general is that they have the ability to reproduce in a variety of different ways, and many plant species are not limited to seeds as their only means of dispersal. A paper by Scott Zona and Cody Coyotee Howard, published in Flora (February 2022), introduces us to the intriguing world of aerial vegetative diaspores – just one of the many ways that plants have to get around.

A diaspore is a plant structure that facilitates dispersal. Seeds are diaspores, as are spores, which are produced by non-seed bearing plants like mosses and ferns. If you’ve ever planted bulbs, you’ve handled another type of diaspore. Bulbs and corms, which many spring flowering plants are grown from, form little offshoots called bulblets and cormels that, when detached from their parent structure, can grow into new individuals. These vegetative diaspores are produced below ground. Aerial vegetative diaspores, on the other hand, are formed on above ground plant parts. This clunky term encompasses a number of different structures that are often simply called bulbils, which Zona and Howard explain is used as “a catch-all term that obscures their morphological identity.”

Compiling a list of plant species that feature aerial vegetative diaspores is a difficult task when plant descriptions from various sources use a broad selection of terminology for the same or similar plant parts. To help complete this task, Zona and Howard defined five distinct types of aerial vegetative diaspores – plantlets, bulbils, cormlets, tubercles, and gemmae – and came up with a list of 252 taxa that are known to feature at least one of these structures.

plantlets on the leaf margin of Kalanchoe daigremontiana (wikimedia commons; Aurélien Mora)

Plantlets are miniature plants attached to another plant. Once mature, they have clearly visible leaves, stems, and roots (or root initials) and are non-dormant, meaning they are ready to grow on their own as soon as they’re given the opportunity. The tiny plants borne along the margins of the leaves of mother of thousands (Kalanchoe daigremontiana) is a great example of a plantlet.

A bulbil consists of a shortened stem surrounded by scale leaves modified for food and water storage. Sometimes root initials are visible at the base of the bulbil. Bulbils remain dormant until they are dispersed and conditions are suitable for growth. When bulbils start growing but remain attached to the plant, they become a plantlet. A good example of a bulbil can be found on bulbous bluegrass (Poa bulbosa).

Cormlets are comprised of stem tissue and, like plantlets and bulbils, have a single axis of polarity. They have highly reduced scale leaves and are dormant at dispersal. Bulbil bugle lily (Watsonia meriana), despite its misleading common name, is a good example of a plant that produces cormlets.

Tubercles are made up of swollen stem tissue and, like tubers (their underground counterparts), have multiple shoot buds and multiple axes of polarity (meaning there is no right side up like there is in plantlets, bulbils, and cormlets). They lack scale leaves and are dormant at dispersal. Air potato (Dioscorea bulbifera) is an example of a tubercle-producing plant. As you might guess from the common name, potato-like structures are produced aerially on this vining plant that was introduced to North America from Africa and is now invasive in Florida.

A gemma is a tiny cluster of undifferentiated cells. Gemmae are non-dormant and lack polarity. They are the smallest and least common form of aerial vegetative diaspore and can be found on Drosera pygmaea, a species of sundew native to parts of Australia and New Zealand.

Drosera pygmaea (wikimedia commons; Björn S…)

Zona and Howard’s list of plants with aerial vegetative diaspores is the most comprehensive list to date – although it is undoubtedly and understandably missing some – and includes representatives from 42 plant families and 21 plant orders. Plantlets are the most common form of aerial vegetative diaspore at 116 taxa, with bulbils coming in second at 72. Cormlets and tubercles are less common, with 25 and 16 taxa respectively. Their paper includes the full list and offers further information about many of the species listed. It’s worth taking time to explore and is a valuable resource for anyone interested in the topic. In addition, their discussion section highlights a number of questions that warrant further investigation.

Questions surrounding reproductive strategies and the dispersal of aerial vegetative diaspores are particularly interesting. Because these structures are vegetative, they are essentially clones of the parent plant, meaning there is no genetic mixing as occurs when seeds are produced. This can be an advantage when sexual reproduction isn’t possible due to lack of pollinators, environmental restrictions, or chromosomal/polyploidy anomalies. It also assures that new individuals are pre-adapted to the site, and it can help a species colonize an area quickly. This ability to rapidly colonize explains why several of the species on Kona and Howard’s list are known to be invasive in parts of the world outside of their native range.

A species that produces both seeds and aerial vegetative diaspores may have an advantage when it comes to dispersal since both types of diaspores have their strengths. Seeds can remain dormant in the soil and are likely to persist in the environment longer than vegetative diaspores, but vegetative diaspores can be produced without relying on pollinators and can establish new individuals quickly. The modes of dispersal between the two can also vary. Seeds can be dispersed by wind or carried away by animals, while vegetative diaspores often rely solely on gravity to get around. One exception is hitchhiker elephant ear (Remusatia vivipara), whose bulbils are equipped with tiny hooks that cling to animal fur and are transported in a similar manner to burs.

hooked bulbils of hitchhiker elephant ear (Remusatia vivipara) (wikimedia commons; Dinesh Valke)

When the advantages of aerial vegetative diaspores are considered, it is a wonder that we don’t see them more often. Many plants can be easily propagated by taking stem, leaf, and/or root cuttings and placing them in conditions that favor adventitious root and shoot growth. This may suggest that dormant genetic pathways for producing vegetative diaspores exist in most plants. Or maybe not. Genetic studies of species that feature these structures are needed in order to understand why they are found in some species and not others. Kona and Howard leave us with a slew of research questions like this, and it’s a topic I’ll continue to check in on.


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Dispersal by Bulbils – A Bulbous Bluegrass Story

The main way that a plant gets from place to place is in the form of a seed. As seeds, plants have the ability to travel miles from home, especially with the assistance of outside forces like wind, water, and animals. They could also simply drop to the ground at the base of their parent plant and stay there. The possibilities are endless, really.

But what about plants that don’t even bother making seeds? How do they get around? In the case of bulbous bluegrass, miniature bulbs produced in place of flowers function exactly like seeds. They are formed in the same location as seeds, reach maturity and drop from the plant just like seed-bearing fruits, and are then dispersed in the same ways that seeds are. They even experience a period of dormancy similar to seeds, in that they lie in wait for months or years until the right environmental conditions “tell” them to sprout. And so, bulbils are basically seeds, but different.

bulbous bluegrass (Poa bulbosa)

Bulbous bluegrass (Poa bulbosa) is a Eurasian native but is widely distributed outside of its native range having been repeatedly spread around by humans both intentionally and accidentally. It’s a short-lived, perennial grass that can reach up to 2 feet tall but is often considerably shorter. Its leaves are similar to other bluegrasses – narrow, flat or slightly rolled, with boat-shaped tips and membranous ligules – yet the plants are easy to distinguish thanks to their bulbous bases and the bulbils that form in their flower heads. Their bulbous bases are actually true bulbs, and bulbous bluegrass is said to be the only grass species that has this trait. Just like other bulb-producing plants, the production of these basal bulbs is one way that bulbous bluegrass propagates itself.

basal bulbs of bulbous bluegrass

Bulbous bluegrass is also propagated by seeds and bulbils. Seeds form, like any other plant species, in the ovary of a pollinated flower. But sometimes bulbous bluegrass doesn’t make flowers, and instead modifies its flower parts to form bulbils in their place. Bulbils are essentially tiny, immature plants that, once separated from their parent plant, can form roots and grow into a full size plant. The drawback is that, unlike with most seeds, no sexual recombination has occurred, and so bulbils are essentially clones of a single parent.

The bulbils of bulbous bluegrass sit atop the glumes (bracts) of a spikelet, which would otherwise consist of multiple florets. They have dark purple bases and long, slender, grass-like tips. Bulbils are a type of pseudovivipary, in that they are little plantlets attached to a parent plant. True vivipary occurs when a seed germinates inside of a fruit while still attached to its parent.

Like seeds, bulbils are small packets of starch and fat, and so they are sought ought by small mammals and birds as a source of food. Ants and small rodents are said to collect and cache the bulbils, which is one way they get dispersed. Otherwise, the bulbils rely mostly on wind to get around. They then lie dormant for as long as 2 or 3 years, awaiting the ideal time to take root.

bulbils of bulbous bluegrass

Bulbous bluegrass was accidentally brought to North America as a contaminant in alfalfa and clover seed. It was also intentionally planted as early as 1907 and has been evaluated repeatedly by the USDA and other organizations for use as a forage crop or turfgrass. It has been used in restoration to stabilize soils and reduce erosion. Despite numerous trials, it has consistently underperformed mainly due to its short growth cycle and long dormancy period. It is one of the first grasses to green up in the spring, but by the start of summer it has often gone completely dormant, limiting its value as forage and making for a pretty pathetic turfgrass. Otherwise, it’s pretty good at propagating itself and persisting in locations where it hasn’t been invited and is now mostly considered a weed – a noxious one at that according to some states. Due to its preference for dry climates, it is found most commonly in western North America.

In its native range, bulbous bluegrass frequently reproduces sexually. In North America, however, sexual reproduction is rare, and bulbils are the most common method of reproduction. Prolific asexual reproduction suggests that bulbous bluegress populations in North America should have low genetic diversity. Researchers set out to examine this by comparing populations found in Washington, Oregon, and Idaho. Their results, published in Northwest Science (1997), showed a surprising amount of genetic variation within and among populations. They concluded that multiple introductions, some sexual reproduction, and the autopolyploidy nature of the species help explain this high level of diversity.

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Interested in learning more about how plants get around? Check out the first issue of our new zine Dispersal Stories.