“How galling!! The audacity of this insect making a home in me!”
True; no matter how one organism looks at it, it’s annoying. The word ‘gall’ originates from Middle English (~ 900 A.D) and refers to bile, the bitter fluid from the gall bladder. The figurative word ‘galling’ refers to irritating, offensive, audacity and very annoying behavior. But how did an abnormal plant growth acquire the same name, gall?
We may never know.
As a child roaming the woods and wild fields, I would often collect tree and shrub leaves and twigs that had protruding bumps in a variety of shapes. I wondered what these odd shapes were, but it never occurred to me that they might be injurious to the plant, or even malicious at all. Nor did I know then how they were formed.
One day while wandering in the field I found a particularly large growth on the stem of a shrub. Pulling out my magic little ‘looking glass’ (pocket magnifier), I watched half a dozen little translucent bugs crawl out of the ball-shaped growth. In a short time, these bugs acquired color and their wings unfolded away from their bodies. I wondered if the abnormal-looking ball of green was a home for these bugs, and only much later did I learn they were called ‘galls’. And from then on, anytime a person uses the word ‘gall’ or ‘galling’, all I can think of are these appropriated plant cells that serve as a home for small insects.
Five decades later and I’m still fascinated by galls!
Here in the high desert of the Great Basin, galls are common on sagebrush, the most dominant plant. What surprises me is the morphological variety of these galls: the colors, shapes and sizes. So, like the child I was (and probably still am), I have been collecting samples to take back with me, as well as photographing them.
What are galls, anyway?
Galls are an abnormal plant growth induced by various parasitic organisms (1), usually insects. These latter galls will be the focus of a series of posts here as I find examples.
Galls serve as ‘incubators’ for developing insects where they gain nutrition and protection from environmental conditions and predators. Some galls are colorful and easily distinguished from the other plant material. Some are wooly, some round and colorful like tiny plums, some are lobed, and others have spiky protuberances.
Gall-inducing insects are usually species-specific and sometimes tissue-specific on the plants they parasitize. Galls can be found on leaves, stems, shoots, flowers and roots. Combined with gall morphology, these traits will often help to identify which insect is associated with them. However, identifying the insects inside will be the confirmation.
These insects manipulate and exploit the chemistry and physiology of plant tissue to their own benefit and development. Accordingly, galls act as physiological sinks for mobilized plant resources, mostly as nutrition for larvae. Fungi sometimes grow on the interior of the gall surface on which the larvae feed.
Like little houses, galls physically serve as protection from the sun, wind, rain and snow. In fact, because the gall-forming insects control gall formation so well, galls are commonly referred to as their extended phenotype. However, several predatory insects have also adapted to this system by inserting their own larvae inside galls. Then a battle for who eats whom ensues until maturation of one or both species. It’s not uncommon to have more than one species of insect emerge from a gall, but only one of those species induces galls.
Protection is one explanation for the high levels of compounds, such as phenolics and tanins, found in many galls. This is considered a defensive gall trait, protecting the gall against natural enemies outside. Thus, in addition to serving as a nutrition sink and physical protection, some galls have a natural chemical defense.
Sagebrush gall midges
Like any plant, it’s an insect-eat-leaf world out there for sagebrush. Of the 237 species of insects that are associates of sagebrush, 42 are gall-forming insects. Of those, the most predominant are Cecidomiidae, or gall midges. They are a small family of tiny flies that are associated with gall induction.
The most abundant gall midges found on sagebrush are of the Rhopalomyia genus. Although there are 32 species, not all may be present in the same location and area. A recent study suggests that land use or local abiotic conditions may greatly influence the diversity of gall midges.
The adult midges lay eggs in the sagebrush stem tissue. The eggs hatch and the larva secrete saliva into the plant. Compounds in the saliva alter the growth of the injured plant cells and the tissue produces a swelling, or gall, around the young insects. However, the size, shape and color of the developing gall are typically specific to the gall midge species. On the other hand, one species unusually induces a wide range of gall morphologies.
During a recent camping trip on Steens Mountain in SE Oregon (and bordering the Refuge), I found many specimens of Medusa galls (Rhopalomyia medusa) on Big Sagebrush (Artemesia tridentate). As seen in the photograph, these galls are composed of numerous leaf-like structures. Looking at the long miniature leafy structures, it’s easy to see how this gall was called “Medusa”.
Medusa Gall on Big sagebrush
The galls develop in October and rest during the winter. They reach full size in the following spring and adult midge flies emerge in April or May. When I was there, May 9-11, the galls were intact with no sign of emergent flies. Considering the elevation (7,300 feet) where I was hiking, patches of snow were common and the climate was barely spring-like.
Authors of a study (2) sampled arthropod diversity on sagebrush in two ecosystems, one surrounded by dryland agriculture and the other area protected from agriculture and significant human use. Their data suggests that diversity of gall midges is highly variable with the dynamics of arthropod-sagebrush interactions and the sagebrush ecosystem. Interestingly, R. medusa was one of a few species that served as an indicator species in low human impact sagebrush habitats. A good description of where I found the many specimens on Steens Mnt.
So, do these galls negatively affect the sagebrush? We will examine that question in a later post!
1. Some bacteria species can also cause galls. This was my first introduction to galls in undergraduate university. Crown gall (Rhizobium radiobacter, formerly known as Agrobacterium tumefaciens) is the textbook and lab example used in plant pathology and lab classes. It is also a common tool to teach Koch’s Postulates. Soil bacterium inserts a small segment of DNA (T-DNA) from a plasmid and into the plant cell. This DNA encodes for genes that produce a plant hormone, auxin (indole-3-acetic acid), via a special pathway that is not used in most plants. Thus the plant has no molecular means of regulating the production of the exocrine hormone. The T-DNA also signals extra production of a group of plant hormones called cytokines, which are involved in cell division. These hormones are responsible for the tumor-like growth of plant tissue and form the galls.
2. Sanford, M.P., Huntly, N.J. 2010. Seasonal patterns of arthropod diversity and abundance on Big sagebrush, Artemisia tridentata. Western North American Naturalist, 70(1): 67-76.