To grow or to flower, that is the cactus conundrum

Melocactus intortus

Flowers are costly structures for plants to produce. In the flowering plant world, there is always a trade-off between growth and reproduction. Flowers are produced from tiny structures called axillary buds, and many plants can only produce one flush of flowers per bud. Cacti are no exception to this rule and their amazing morphological adaptations to harsh climates has forced them into quite a conundrum when it comes to reproduction.

The axillary buds of cacti are located at the base of their spines in little structures called areoles. This is where the flowers will eventually emerge. However, unlike plants that can produce cheap stems and branches, cacti must produce a whole new chunk of stem or internode before they can produce more axillary buds. Think of it this way, if a cactus wants to produce 10 flowers, it must produce ten internodes to do so. This means producing all of the expensive cortex and epidermis along with it. Their harsh environments have forced most cacti into an extremely tight relationship between growth, water storage, photosynthesis, and flowering that is potentially very limiting from a reproductive standpoint.

Micranthocereus estevesii with lateral cephalium

Amazingly, some cacti have managed to break from this evolutionary relationship and they have done so in a bizarre way. Take a look at all of the cacti pictured here. Each has developed a strange looking structure called a cephalium. Essentially, you can think of the cephalium of a cactus as its “adult” reproductive form whereas the rest of the body consists of non-reproductive, photosynthetic “juvenile” form.

The cephalium is a unique and fascinating structure. It differs from the rest of the cactus body in that it is not photosynthetic. It also produces no chlorophyll and no stomata. In fact, it does not form anything like the epidermis of the rest of the plant. Instead, the cephalium produces dense clusters of short spines and trichomes. Most importantly, it produces tightly packed axillary buds in high abundance. These are the buds that will produce the flowers. The end result is a wacky looking structure that has the ability to produce far more flowers than that of cacti that do not grow a cephalium.

Facheiroa tenebrosa with lateral cephalium

Obviously not all cacti produce cephalia but it is common in genera such as Melocactus, Backebergia, Espostoa, Discocactus, and Facheiroa (this is not a complete list). What the cephalium has done for genera like these is decouple the afore mentioned relationships between growth and reproduction. For a period of time (often many years) following germination, these cacti grow the typical succulent, photosynthetic stems we are accustomed to seeing.

At some point in their development, something triggers these plants to switch to their adult forms. Axillary buds within either lateral or apical meristems switch their growth habit and begin forming the cephalium. It is worth mentioning that no one yet knows what triggers this switch. If the cephalium is produced from axillary buds in the apical meristem like we see in Melocactus, the plant will no longer produce photosynthetic tissues. This represents another major trade-off for these cacti. Such species must rely on the photosynthetic juvenile tissues for all of their photosynthetic needs for the rest of their lives (unless the cephalium is damaged or lost). Backebergia have managed to get around this trade-off by not only growing multiple stems, they will also shed their apical cephalia after a few years, thus re-initiating photosynthetic juvenile growth.

Backebergia militaris with bizarre apical cephalia reminiscent of the bearskin hats of the Queen’s guard.

Things are a bit different for cacti that produce lateral cephalia. Genera such as Espostoa, Facheiroa, and Buiningia are less limited by their cephalia because they are produced along the ribs of the stem, thus leaving the apical meristem free to continue more typical photosynthetic growth. Nonetheless, the process is much the same. Dense clusters of spines, trichomes, and most importantly, axillary buds are produced along the rib, giving each stem a lovely, lopsided appearance.

There are other benefits to growing cephalia in addition to simply being able to produce more flowers. The densely packed spines and trichomes offer the developing flowers and fruits ample protection from both the elements and herbivores. Floral buds are free to develop deep within the interior of the cephalium until they are mature. At that point, the cells will begin to swell with water, pushing the flower outward from the cephalium where it will be exposed to pollinators. As the petals curl back, they offer a safe spot for visiting pollinators that is free from menacing spines. Once pollination has been achieved, the flower wilts and the deeply inferior ovaries are then free to develop within the safety of the cephalium. Once the fruits are mature, they too will begin to swell with water and be pushed out from the cephalium where they will attract potential seed dispersers.

Melocactus violaceus with fruits emerging from the cephalium

I hope that I have convinced you of just how awesome this growth form can be. I will never forget the first time I saw a cactus topped with a cephalium. It was a mature Melocactus growing in a cactus house. Sticking out of the odd “cap” on top was a ring of bright pink fruits. I knew nothing of the structure at that time but it was incredible to see. Now that I know what it is and how it functions, I am all the more appreciative of these cacti.

This post was inspired by the diligent work of Dr. Jim Mauseth. Click here to learn more about cacti.

Photo Credits: [1] [2] [3] [4] [5]

Further Reading: [1] [2] [3]

Himalayan snowball plants and their fashionably functional coats

Credit to CGTN Nature film crew

Hairy plants are both fun and functional. Hairs or trichomes on the leaves of plants can serve a variety of functions. If the plant is growing in a region prone to cold temperatures, it is thought that a dense layer of hairs can function like a wool coat, keeping the plant warm when temperatures drop. This is such a popular idea that it is often assumed rather than tested. For a strange group commonly referred to as Himalayan snowball plants, the truth is a bit more complicated.

Himalayan snowball plants are members of the genus Saussurea, which hails from the family Asteraceae. Though the genus is widespread, the Himalayan snowball plants are confined to high elevation, alpine habitats in central Asia. As you can imagine, life at such altitudes is defined by extremes. Temperatures during the day can skyrocket due to the lack of atmospheric insulation. Conversely, temperatures can take a dive as weather changes and/or the sun goes down. One look at the Himalayan snowball plants tells you that these plants are wonderfully adapted to such habitats. But what kind of advantages does that this coat of hair provide?

Credit to CGTN Nature film crew

Well, research has revealed a bit more nuance to the whole “winter coat” idea. Indeed, it does appear that the furry coat does in fact provide some insulation to the plant. However, most of the warmth appears to come from the dark color of the inflorescence rather than by pure insulation alone. After all, the vast majority of plants do not produce any heat. The flower heads or capitula of these daisy relatives is low in stature. This keeps it out of the way of the coldest winds. Also, they are so deeply violet in color that they can appear black. This is no accident. As anyone can tell you, darker colors absorb more heat and that is exactly what happens with the Himalayan snowball plants.

Another interesting thing to consider is that most of the growth and reproduction in these plants occurs during frost-free periods of the year. Though temperature swings are frequent, it rarely gets cold enough to severely damage plant tissues until long after the plants have flowered and set seed. Moreover, there is some evidence to suggest that the dense coat of hairs may have a cooling effect during periods of intense exposure to sunlight. Their light color may reflect a lot of the incoming radiation, sparing the plant from overheating. Therefore, it appears that the benefit of such a thick coat of hairs has more to do with avoiding temperature swings than it does ensuring constant warmth. By buffering the plant against huge swings in ambient temperature, the hairs are able to maintain more favorable conditions for plant growth and reproduction.

Credit to CGTN Nature film crew

Also, because this area experiences a monsoon season during growth and flowering of Himalayan snowball plants, these hairs may also serve to repel water, keeping the plants from becoming completely saturated. If water were to stick around for too long, it could open the plant up to pathogens like fungi and bacteria. It could also be that by insulating the plant against temperature swings, the hairs also provide a more favorable microclimate for pollinators. Bumblebees are thought to be the main pollinators of Himalayan snowball plants and despite their ability to maintain higher internal temperatures relative to their surroundings, anything that can buffer them as they feed would be beneficial to both the bees and whatever plant they may be pollinating as a result.

Photo Credit: [1]

Further Reading: [1] [2]

How Air Plants Drink

 Tillandsia tectorum. Photo by Edu licensed under CC BY-NC-ND 2.0

Air plants (genus Tillandsia) are remarkable organisms. All it takes is seeing one in person to understand why they have achieved rock star status in the horticulture trade. Unlike what we think of as a "traditional" plant lifestyle, most species of air plants live a life free of soil. Instead, they attach themselves to the limbs and trunks of trees as well as a plethora of other surfaces. 

Living this way imposes some serious challenges. The biggest of these is the acquisition of water. Although air plants are fully capable of developing roots, these organs don't live very long and they are largely incapable of absorbing anything from the surrounding environment. The sole purpose of air plant roots is to anchor them to whatever they are growing on. How then do these plants function? How do they obtain water and nutrients? The answer to this lies in tiny structures called trichomes. 

Trichomes are what gives most air plants their silvery sheen. To fully appreciate how these marvelous structures work, one needs some serious magnification. A close inspection would reveal hollow, nail-shaped structures attached to the plant by a stem. Instead of absorbing water directly through the leaf tissues, these trichomes mediate the process and, in doing so, prevent the plant from losing more water than it gains. 

The trichomes themselves start off as living tissue. During development, however, they undergo programmed cell death, leaving them hollow. When any amount of moisture comes into contact with these trichomes, they immediately absorb that water, swelling up in the process. As they swell, they are stretched out flat along the surface of the leaf. This creates a tiny film of water between the trichomes and the rest of the leaf, which only facilitates the absorption of more water. 

Trichomes up close.  Photo by Mark Smith1989 licensed under CC BY-SA 4.0

Because the trichomes form a sort of conduit to the inside of the leaf, water and any nutrients dissolved within are free to move into the plant until the reach the spongy mesophyll cells inside. In this way, air plants get all of their water needs from precipitation and fog. Not all air plants have the same amount of trichomes either. In fact, trichome density can tell you a lot about the kind of environment a particular air plant calls home. 

Photo by Bernard DUPONT licensed under CC BY-SA 2.0

The fuzzier the plant looks, the drier the habitat it can tolerate. Take, for instance, one of the fuzziest air plants - Tillandsia tectorum. This species hails from extremely arid environments in the high elevation regions of Ecuador and Peru. This species mainly relies on passing clouds and fog for its moisture needs and thus requires lots of surface area to collect said water. Now contrast that with a species like Tillandsia bulbosa, which appears to have almost no trichome cover. This smoother looking species is native to humid low-land habitats where high humidity and frequent rain provide plenty of opportunities for a drink. 

Photo by Bocabroms licensed under CC BY-SA 3.0

Absorbing water in this way would appear to have opened up a plethora of habitats for the genus Tillandsia. Air plants are tenacious plants and worthy of our admiration. One could learn a lot from their water savvy ways. 

Photo Credits: [1] [2] [3] [4]

Further Reading: [1] [2] [3]