Allelopathy Helps Black Walnuts Compete

A walk through the woods can be one of the most peaceful and calming experiences — a place where you can find quiet for reflection and marvel at the beauty of nature. Little do most people know that some plants, especially one specific tree, wage chemical warfare against other plants to keep away potential neighbors that would compete for nutrients and sunlight. In the Appalachian Mountains, the tree most skilled at chemical warfare is the black walnut.

The black walnut tree (Juglans nigra) is a useful, yet often misunderstood tree. Prized for its excellent wood qualities for lumber and furniture, the nuts it produces are either loved or reviled by those who try them.

The flavor of black walnuts is hard to describe. I would say that they have an almost astringent flavor, mainly due to the high level of tannins in them. They aren’t my favorite, but I don’t mind them either. I’ve learned to accept them, unlike during my childhood when you knew which church lady’s cake to avoid at the potluck because you knew that she put black walnuts in everything she baked.

My appreciation for black walnuts grew the year that I was the official nut judge (no joke) for the Black Walnut Festival in Spencer, WV. It was quite an experience — examining and weighing all the entries with a team of high school FFA students who cracked more than a few inappropriate jokes about the situation.

You could tell when someone was picking or cracking black walnuts, thanks to the tannin stains on their hands that just wouldn’t wash off. Black walnuts are a tough nut to crack (literally), so I also remember my grandmother cracking them “the easy way.” She would just pile them up in the driveway and run over them a time or two with her behemoth of an Oldsmobile (you know, the one that had full seats front and back and could hold half the neighborhood).

Black walnut trees have the interesting ability to excrete a chemical called juglone, which makes it nearly impossible for a number of plants to grow anywhere in its root zone. Juglone works by damaging the tiny root hairs on roots that are responsible for taking up a great majority of the water and nutrients the plants use. Research shows that it also interferes with the interaction of the roots with mycorrhizal fungi that aid the plant in taking up nutrients.

This process is not just specific to black walnuts. There are several other plants that do this. The phenomenon, called allelopathy, occurs when an organism excretes something that inhibits the growth of other things around it. You could equate it to the Penicillium fungus excreting a chemical that kills bacteria around it. We harness that chemical to use as penicillin.

Some plants are especially sensitive to the chemical. Many vegetable plants, especially tomatoes, are sensitive. Some plants, mainly those that would grow wild in the woods, are not susceptible. Many grasses also have a hard time growing beneath black walnut trees (tall fescue and Kentucky bluegrass being the exception, except during periods of drought).

Publication with lists of plants tolerant and damaged by juglone

All parts of the tree produce the juglone chemical, so the effects could spread beyond the perimeter of the tree from fallen leaves and branches. I would also suggest that you make sure any fresh woodchip mulch that you use (specifically that from local tree cutters) is free of black walnut. The juglone may break down after composting the wood chips for six months to a year, but I would still be cautious about its use. The wood will release the chemical, killing susceptible plants for a few years in the area where it is applied. Studies suggest that juglone will break down during the composting process, but I would check to make sure by starting a few tomato seeds on the batch of compost to see what happens.

—Garden Professor John Porter is a county extension agent for West Virginia University and writes the weekly Sunday garden column for the Charleston Gazette-Mail Newspaper.  This article was originally published October 2, 2015.

You can find John’s writing at and on Facebook and Twitter.

“Lazy” corn and gravitropism

Inspired by Linda’s post about thigmomorphogenesis, I decided today I would add the word gravitropism to your vocabulary. It simply means growth in response to gravity. Shoots of plants grow up, because they are negatively gravitropic, they grow against the pull of gravity, while roots are positively gravitropic and grow down towards the pull of gravity.

And why is that so important? Well… this is what happens when gravitropism is missing.


To the left is normal old corn. The plant to the right was not sat on by a raccoon or anything, it simply has a mutation in a gene called lazy plant1. I’m not kidding. That’s the official, scientific name for this gene. Geneticists have fun with their names, though fruit fly geneticists are for sure the kings of silly gene names. This gene got that name because, as you can see, without a functioning copy of that gene, the corn plant no longer can detect the pull of gravity and so flops down in a “lazy” manner.

This corn is just odd, of course, with no real value (though it was fun to grow) but similar mutations are what give us some of the “weeping” or trailing forms of popular ornamental trees and shrubs.

Your new word for the day: thigmomorphogenesis

I just finished reviewing 4 manuscripts for three different journals and boy is my brain fried. My private reactions ranged from “I can’t wait until this one is published!” to “If I were to use sheet mulch this manuscript would be my first choice.” Anyway, it was the latter manuscript that got me to thinking about what can go wrong with experimental design, which brings up today’s word: thigmomorphogenesis.

This is a great word for those who enjoy figuring out word meanings by deciphering the (usually) Greek or Latin roots. (This exercise also helps you figure out how to pronounce it.) We have “thigmo-” which means touch, “-morpho-” which means appearance, and “-genesis” which means beginning. String them all together and you get the phenomenon seen when plants respond to mechanical stimulation by changing their growth pattern and hence the way they look.

Wind direction from the right creates an asymmetric hedge.
Wind direction from the right creates an asymmetric hedge.

You can easily see examples of thigmomorphogenesis in everyday life. Look at a line of hedge plants where the plants on the end are more susceptible to wind movement and brushing by people, animals or vehicles. They are always shorter, aren’t they? Plants subjected to chronic thigmomorphogenic forces are generally shorter than their neighbors and thicker in girth. (For a longer discussion about how thigmorphogenesis works, you can read my online column.)

How does all of this relate to experimental design? Well, think about what happens if you are testing a product that requires applying it to the leaves of plants once a week. Your treatment plants are touched every week. How can you know that any changes in your experimental plants aren’t due to being touched? The way you eliminate this source of variability is by treating all of the plants the same way. When you are applying the product to the treatment leaves, you apply water (or whatever the solvent is for the product in question) to the control leaves. That way thigmomorphogenesis remains just an interesting tongue-twister and not a fatal design flaw in an experiment.

Puya report!

For all five of you that might have paid attention to my posts on the genus Puya (which does in fact rhyme with booyah…thank you my west-coastie friends):

Here’s the update that you’ve been waiting for!

Puya is a horrifically spiny, painful, and hateful genus in the Bromeliad family. Native to the Andes, the fish-hook-like spines snare passing mammals; the rotting flesh provides nutrients to the exceptionally lean soil of the arid steppes on which it sort of grows/becomes grumpier.

Puya flowers once an eon, in a spectacular [but ill-earned] display that turned me to mush, based on a photo in an Annie’s Annuals catalog (see my “eternal gardening optimist” post). Autumn of 2012, I ordered and received one healthy Puya berteroniana in a 4” pot. Heckling commenced.  Overwinters in a 40 F greenhouse, where it was watered once or twice. Summers have been spent on our deck. Osmocote has hopefully provided required nutrients. Expected to kill her within months, as it is SO VERY not native to the verdant and humid Blue Ridge mountains of Southwest Virginia.

Happy and amazed to say Pootie [what was I going to name her? Bert??] is in her 3rd year – continuing to grow, and, AND, captured her very first mammal!


Okay… so it’s a fluffy stuffed possum, and the dogs dropped it from the deck above. But snagged! You know Pootie got a thrill…

When Plants Attack! (each other)

When you talk about killer plants, your mind may conjure images of a man-eating plant in “Little Shop of Horrors,” insect-eating Venus flytraps or poisonous plants like deadly nightshade.

While all of those scenarios are interesting in and of themselves, what about plants that attack other plants?

I’m talking, of course, about parasitic plants. These plants thrive on stealing nutrients from other plants, either weakening them or, quite possibly, killing them.

Parasitic plants connect themselves to a host plant and siphon off the sugars that plant produces and the nutrients it pulls from the soil. These plants often bend the definition we have in our heads of plants, since they don’t have to behave like other plants that make their own food.

Probably the most well-known (and beloved) parasitic plant is mistletoe. The plant that gives us the warm fuzzies and romantic feelings around the holidays makes its living by feeding off of the trees in which it lives. They don’t talk about that aspect of the plant in all those Christmas songs. It doesn’t kill the tree, but a heavy infestation can weaken a tree and slow its growth.

Indian pipe (Monotropa uniflora). Photo courtesy GP Raymond Eckhart

While they are few in number, there are some parasitic plants you may run into. Another parasitic plant in our part of the world is the Indian pipe (Monotropa uniflora), a white, chlorophyll-free plant that resembles a smoking pipe as it unfurls from the forest floor. Without chlorophyll, it can’t make its own food, so it connects itself to a nearby tree (usually beech) for nutrients.

Another plant, called a beech drop (Epifagus americana), also makes its living in the same manner. A plant called squaw root or bear corn (Conopholis americana), because it resembles an ear of corn growing out of the forest floor, is a parasitic plant that connects with the roots of oak trees.

An infestation of dodder beginning in an annual bed. Photo courtesy Ann Berry.
An infestation of dodder beginning in an annual bed. Photo courtesy Ann Berry.

These plants may cause a little damage to their host plants. This week, though, there seems to be something more sinister afoot. I received two different calls about the same parasitic plant this week, from different parts of West Virginia (one of which came from Ann Berry, associate vice president for marketing and outreach at WVU). It seems that the problem here was with a parasitic plant called dodder (Cuscuta sp.). Despite the name, I assure you that this plant does not dodder around when it comes to feeding off other plants. This plant can severely infect and potentially kill any plant it touches.


Seeds of the plant germinate in the soil, so it starts life just like any other plant. Once germinated, though, the seedling has about 10 days to find a host plant to attach to and begin feeding. But this is not left to chance — it seems that dodder is a pretty good hunter. Scientists have determined that dodder can, in a way, sense chemical signals from nearby plants and grow directly toward them.

Dodder is an odd-looking plant, and many people don’t even know to classify it as a plant. It grows in long strings, often without leaves (or only having inconspicuous ones). Different species can be different colors. The one that is most common here is often a yellow-orange color.

cuscuta Haustorium
Dodder, above, inserts a haustorium into its host plant.

Once the dodder touches the soft tissue of a plant (leaves or stems), it inserts a structure called a haustorium into the plant. Haustoria insert themselves into the plants vascular tissue (veins) and siphons off the water, sugars and nutrients. After the connection is made, the dodder plant detaches its roots from the ground and becomes completely reliant upon the host plant. Luckily it has trouble attacking woody plants, so it mainly goes after herbaceous ones.

One connection is bad enough, but the dodder twines its way around the plant as it grows, resembling what some would call “silly string.” Everywhere the dodder touches the host, it sends in new haustoria to strengthen its connection. If other plants are close enough, the dodder will grow outward through the air to ensnare another host. It can easily grow to encompass many plants, covering them completely and eventually strangling them or starving them out.

My advice to both of the callers this week was to remove as much of the plant as possible, as soon as possible. Unfortunately, the plant can regrow from the connections it makes with the host plant, so you often need to remove whole parts of the plant or the whole plant itself. If it has only made one or two connections, you may be able to control it just by removing the dodder from the plant.

Dodder is hard to see on the ground as it germinates, so it is only usually spotted after it has attached and grown on a plant. If you do happen to catch it before it attaches to a plant, cultivating the soil to break it up and removing as much by hand as possible will help. Unfortunately, there is no spray or control method that will kill the dodder without killing the host.

Dodder is definitely a bizarre plant that many have not seen. Keep an eye out for it this year, since it seems to be cropping up in unexpected places. It just goes to show you that sometimes it’s a plant-eat-plant world out there.

This article was originally published 08.09.15 in the Charleston Gazette-Mail.  You can find more article at

When half a flower is the wrong color…

If you’ve spent enough time around flowers, you’ve probably seen this. It isn’t exactly common, but it happens, and is so distinctive that you’ll almost always notice when it happens, as I did on one of my gladiolus the other day.


Everything else is as normal, but a chunk of the flower is white instead of the usual soft peachy white.

What we have here is a sectorial chimera. Chimera means an organism with two (or more, I suppose) genetically different cell types, and a sectorial chimera is when there is one distinct section of the plant made up of a different cell type.

And why is this showing up in my usually pink gladiolus? Well, somewhere early in the development of this flower spike, there was a chance mutation in a cell. That mutation stopped those cells from producing the usual pink pigment, so the mutant cells make white flowers. The new mutation and the original cells continued to grow and divide, so some of the flower is from the newly mutated white form, and some is the original cell type.

Now, when people hear the word “mutant” they either think x-men or nuclear fallout, but the fact is mutations are a perfectly common, normal part of everyday life for organisms, and of course are critically important to continuing evolution.

This type of bicolored flower is cool looking, but certainly a one-off. Sectorial chimeras are very unstable. Next year, most likely, the flowers will just be pink again, or possible a branch will send off pure white flowers. So when you see a sectorial chimera in the garden, take a picture, put it on facebook, and enjoy it because it probably isn’t coming back.

Robbers in the garden

Salvia azurea (maybe my favorite salvia in the world — sky blue flowers in late summer/fall, hardy to zone 5) is blooming in the garden, and the bees are all over the flowers. But while some are poking their heads into the flowers to drink nectar and transferring pollen as they do so, others are up to something more sinister.

This carpenter bee looks like it is pollinating. It isn't.
This carpenter bee looks like it is pollinating. It isn’t.

The evidence of what they are up to is clear if you look closely at the side of the flowers after they leave.

The little hole at the base of the flower is clear sign of an illegal break in.
The little hole at the base of the flower is clear sign of an illegal break in.

See the little hole in the base of the flower? That is where the carpenter bee bit a hole in the flower to get access to the nectar instead of going in the front of the flower as one would expect.

This phenomenon is called nectar robbing because it is an evolutionary betrayal of sorts. Flowers have evolved nectar to lure bees and other pollinators into the flower, so the bees will pollinate while getting their sugar fix. When the bees nectar rob, they’re getting the payment without doing the actual pollinating.

So why bite a hole in the side of the flower instead of just going in the front? Well, many flowers have evolved flower forms that make the nectar hard to reach by anything but their preferred pollinator, the species that most effectively moves pollen from plant to plant. In the case of this salvia, the nectar is down at the base of the flower, and only accessible to bees with long tongues, like whatever species normally pollinates it in its native range in the US plains. In other words, it is out of reach of the carpenter bees to save it for another bee, probably a bumble bee. Which works great to avoid wasting nectar on sub-optimal pollinators… unless, of course, those bees become robbers.

So next time you see bees on your flowers, take a look… they might just be robbers, not pollinators.

Pollination strategies can lead to allergy woes

This article was originally published in my weekly newspaper column in the Charleston Gazette-Mail.  Articles are archived at

A quick selfie with President Gee.  He's perfected the art of the selfie.
A quick selfie with President Gee. He’s perfected the art of the selfie.

A few weeks ago I made my way to South Dakota for the annual meeting of the National Association of County Agricultural Agents (where fellow GP and I made the rounds at the trade show scrutinizing wacky products). It is a fun conference made even more special this year by the fact that WVU President E. Gordon Gee was in attendance as the conference co-keynote speaker and recipient of the Service to American/World Agriculture award.  But I digress…..

Two days into the conference something wasn’t quite right. I kept feeling worse and worse, and by Wednesday I was confined to my hotel room (save for a venture out to the conference banquet for dinner). I would not have been functional for the rest of the trip save for the kindness of a co-worker who went through the pharmacy red tape to procure and deliver “the good stuff” to my hotel room.

I thought I had a sinus infection at best (I get them often) and the flu at worst (yes, it was really that bad). But guess what — I’m just really allergic to South Dakota. Two days after my return, I was nearly back to normal (well, my normal, anyway).

Those who know me know that I suffer from the occupational hazard of allergies. Irony dictates that my allergies are only to about two dozen plants and two molds (that occur in mulch/compost).  Lucky me!

Lots of corn surrounding my hotel in the outskirts of Sioux Falls.
Lots of corn surrounding the outskirts of Sioux Falls.

My best guess is that I had a reaction to the corn pollen of South Dakota. It makes sense — while we do grow some corn here in West Virginia, the Mount Rushmore State boasts an estimated 4.75 million acres of corn. I don’t think I was tested for corn pollen allergies, but since corn is not a major crop here, it may not be part of the common test.


I tell this story not for sympathy (well, OK, maybe a little) but it brings up a good illustration about pollination strategies of plants.

Scanning electron microscope image of pollen grains from a variety of common plants: sunflower (Helianthus annuus), morning glory (Ipomoea purpurea), prairie hollyhock (Sidalcea malviflora), oriental lily (Lilium auratum), evening primrose (Oenothera fruticosa), and castor bean (Ricinus communis).
Scanning electron microscope image of pollen grains from a variety of common plants: sunflower (Helianthus annuus), morning glory (Ipomoea purpurea), prairie hollyhock (Sidalcea malviflora), oriental lily (Lilium auratum), evening primrose (Oenothera fruticosa), and castor bean (Ricinus communis).

You see, plants like corn rely on chance and wind to spread their genes around. In corn, the pollen drops from the male flowers (the tassel on the top) to the stigma of the female flower (the end of the silk sticking out of the cob). The process relies on lots of pollen being released into the air, since there is a good chance that a lot of it will miss the target. Corn pollen is usually heavy, therefore it doesn’t blow too far from the plant (unless there is lots of wind).

This is why you don’t get a good corn crop if you don’t have a big block of corn in the garden — just one or two rows doesn’t drop enough pollen to pollinate all the flowers. When the silks don’t get pollinated, you’ll end up with incomplete cobs missing kernels. This can also happen if the corn is in bloom during a long period of rain — the rain washes all of the pollen off before pollination can occur.

Most of the major allergen-producing plants are wind pollinators — trees, grasses, ragweed. They all release copious amounts of pollen into the air hoping for it to land in the right place.

Some plants still rely on pollen getting moved from plant to plant or flower to flower, but they remove the chance involved with wind pollination. These plants have a stickier pollen that stays on the flower and waits for something to come along and move it — a bee, a butterfly, a moth, a hummingbird, etc. These plants hold on to their pollen and have the more directed approach of getting a courier to make a direct delivery of their pollen between flowers.

Since these plants don’t leave the pollination to chance, they generally produce less pollen. Some good examples are fruit trees (apples, peaches, pears), sunflowers, squash, goldenrod and roses. Since they don’t release it into the air, they usually aren’t considered major allergens.

Still yet, some plants want to take no chance with their next generation. Self-pollinating plants don’t rely on pollen being spread to different flowers — they take care of business themselves. These plants are perfectly fine without crossbreeding with other plants.

Sometimes, these plants are so dedicated to self-fertilization that they make it difficult for the pollen to leave the flower. Bean flowers have a lower lip that curves upward to protect the reproductive parts inside. Tomato flowers are nearly completely enclosed. You may see bees going from flower to flower, but their search for food is in vain — they can’t get into the flower. Their buzzing does help dislodge the pollen inside the flower, but they don’t have access to spread it around. Producers that grow tomatoes in greenhouses where there is no wind to knock the pollen loose either buy boxes of bumblebees to release in the greenhouse, or use something like a vibrating toothbrush to help the flowers self-pollinate (no joke).

This is why you can plant two different tomatoes just a few feet apart and not have them crossbreed, but you would have to plant squash up to two miles apart (or protect the flowers) to guarantee that you get the same variety if you plan on saving seeds. This is why the most commonly saved seeds, at least in this area, are tomatoes and beans — they are easy to guarantee that you won’t get something other than what you plant.

So if you learn anything from this article, check out how plants pollinate before you save their seeds, and take plenty of allergy meds with you if you go to South Dakota.

The cardboard controversy

I’m not a fan of using corrugated cardboard as a mulch, which like other sheet mulches creates problems for the underlying soil. Long-time readers of this blog may remember several previous posts (1, 2, 3 and 4) on this topic and I won’t belabor the points made in those posts. Instead, today I’m doing to focus on cardboard itself.

Cardboard mulch under wood chips
Cardboard mulch under wood chips

First, cardboard is a generic term that can refer to many types of manufactured paper. The box you see delivered to your front door is more properly called corrugated board or containerboard. It consists of two layers of linerboard sandwiching a layer of accordion-like fluting material. The linerboard is made from sheets of pulp that may be coated to improve smoothness (more about this later). The finished linerboard is laminated using adhesives to both sides of the fluting material.

Corrugated boxes are built to be tough.
Corrugated boxes are built tough

These boxes are made to withstand rough handling and to protect the contents from the external environment. It’s tough stuff: while you might be able to bend a piece of corrugated board fairly easily, it’s more difficult to tear it in half. The more heavy duty the box, the more difficult it is to bend or tear its walls.

So let’s now consider using this tough material in your garden as a mulch. It may be coated as mentioned earlier to improve smoothness. That’s going to prevent it from absorbing moisture. The coating also reduces the ability for gases to move between the soil and the atmosphere. In fact, smoothness is measured using an air leak method – the smoothest materials have the least air leakage.

Photo credit vizpix at Flickr
Photo credit: vizpix at Flickr

A garden or landscape mulched with cardboard (or heaven forbid several layers of cardboard as part of the science-free lasagna mulch method) is now covered with a tough, relatively gas- and water-impermeable material that will take some time to break down. It’s hardly a mulch that’s going to nurture soil life.

But cardboard mulch fans swear that they find more earthworms under cardboard than anywhere else in their garden. This is almost always the first response I get from gardeners who don’t believe that cardboard causes problems. And this is where it’s important to consider earthworm behavior.

Photo credit: Kurt B. on Flicker
Photo credit: Kurt B. on Flickr

We’ve all observed that earthworms crawl to the soil surface during heavy rains; this is due in part to water filling their burrows and reducing oxygen availability (Chuang and Chen demonstrated this nicely in 2008). Likewise, the reduction in oxygen movement from the atmosphere into cardboard-covered soil would cause worms to crawl upwards in an effort to find oxygen at the soil surface.

So don’t assume your lasagna mulching draws earthworms to your garden. It’s more likely that you’re smothering their habitat.

***An update on cardboard gas permeability. We’ve just published an article comparing diffusion rates of different mulches. You can find the article here but it is behind a paywall. Here is a graphic comparing diffusion rates of various mulches. This is a logarithmic scale.
Now, until cardboard proponents publish evidence to the contrary, it’s pretty obvious that cardboard mulch interferes with gas diffusion.

***And another update on how our blog works. This post, by far, is the most popular. It generates a lot of comments. All comments must be approved before they’re posted, and  I don’t approve comments that are derogatory or promote a belief in the absense of supporting science. If you want your comment to be published, be polite and provide evidence to support your statements. Otherwise, you are wasting your time.

***Another update on cardboard in your garden: A recent paper reports on PFAs (aka “forever chemicals”) in various products used for poultry bedding (among other things). Cardboard was one of the worst. The article is behind a paywall but I have access to it and was able to find the table shown here. So if you need yet another argument to NOT use cardboard as a mulch (like in “lasagna gardening), maybe “keep forever chemicals out of your garden” will do it.