Lettuce Gone Wild, part two

The lettuce we eat is a close relative to the lettuce we weed out of our gardens. Last week we discussed the potential that wild relatives may have for improving cultivated lettuce. But if wild lettuce can be crossed with cultivated lettuce to create new cultivars, can cultivated lettuce cross with wild lettuce to make it more weedy?

Because so many of our crops are closely related to some of the weeds found along with them or the plants growing in nearby natural areas, the creation of crop-wild hybrids has long been a concern. This concern is heightened in the age of transgenic crops (also known as GMOs), for fear that hybrids between weeds and such crops could create super weeds – fast spreading or highly adapted weeds resistant to traditional control methods such as certain herbicides. To reduce this risk, extensive research is necessary before such crops are released for commercial use.

flowers of prickly lettuce (Lactuca serriola)

There are no commercially available, genetically modified varieties of cultivated lettuce, so this is not a concern when it comes to crop-wild hybrids; however, due to how prevalent weedy species like prickly lettuce (Lactuca serriola) are, hybridization with cultivated lettuce is still a concern. So, it is important to understand what the consequences might be when hybridization occurs.

In a paper published in Journal of Applied Ecology in 2005, Hooftman et al. examined a group of second-generation hybrids (L. sativa x L. serriola), and found that the hybrids behaved and appeared very similarly to non-hybrid prickly lettuce. They also found that the seeds produced by the hybrids had a significantly higher germination rate than non-hybrid plants. This is an example of hybrid vigor. Thus, “if hybridization does occur, this could lead to better performing and thus potentially more invasive (hybrid) genotypes.” However, the authors cautioned that “better performing genotypes do not automatically result in higher invasiveness,” and that much depends on the conditions they are found in, the level of human disturbance, etc.

Another thing to consider is that hybrids are not stable. In an article published in Nature Reviews Genetics in 2003, Stewart et al. adress the “misunderstanding that can arise through the confusion of hybridization and … introgression.” It is wrong to assume that hybrids between crops and wild relatives will automatically lead to super weeds. For this to occur, repeated crosses with parental lines (also known as backcrossing) must occur, and “backcross generations to the wild relative must progress to the point at which the transgene [or other gene(s) in question] is incorporated into the genome of the wild relative.” That is what is meant by “introgression.” This may happen quickly or over many generations or it may never happen at all. Each case is different.

prickly leaf of prickly lettuce (Lactuca serriola)

In a paper published in Journal of Applied Ecology in 2007, Hooftman et al. observe the breakdown of crop-wild lettuce hybrids. They note that “fitness surplus through [hybrid vigor] will often be reduced over few generations,” which is what was seen in the hybrids they observed. One possible reason why this occurs is that lettuce is predominantly a self-crossing species; outcrossing is rare, occurring 1 – 5% of the time thanks to pollinating insects. But that doesn’t mean that a stable, aggressive genotype could never develop. Again, much depends on environmental conditions, as well as rates of outcrossing and other factors relating to population dynamics.

A significant expansion of prickly lettuce across parts of Europe led some to hypothesize that crop-wild hybrids were partly to blame. In a paper published in Molecular Ecology in 2012 Uwimana et al. ran population genetic analyses on extensive data sets to determine the role that hybridization had in the expansion. They concluded that, at a level of only 7% in wild habitats, crop-wild hybrids were not having a significant impact. They observed greater fitness in the hybrids, as has been observed in other studies (including the one above), but they acknowledged the instability of hybrids, especially in self-pollinating annuals like lettuce.

seed head of prickly lettuce (Lactuca serriola)

It is more likely that the expansion of prickly lettuce in Europe is due to “the expansion of favorable habitat as a result of climate warming and anthropogenic habitat disturbance and to seed dispersal because of transportation of goods.” Uwimana et al. did warn, however, that “the occurrence of 7% crop-wild hybrids among natural L. serriola populations is relatively high [for a predominantly self-pollinating species] and reveals a potential [for] transgene movement from crop to wild relatives [in] self-pollinating crops.”


Lettuce Gone Wild, part one

Lettuce, domesticated about six thousand years ago in a region referred to as the Fertile Crescent, bears little resemblance to its wild ancestors. Hundreds of years of cultivation and artificial selection eliminated spines from the leaves, reduced the latex content and bitter flavor, shortened stem internodes for a more compact, leafy plant, and increased seed size, among several other things. The resulting plant even has a different name, Lactuca sativa (in Latin, sativa means cultivated). However, cultivated lettuce remains closely related to its progenitors, with whom it can cross to produce wild-domestic hybrids. For this reason, there is great interest in the wild relatives of lettuce and the beneficial traits they offer.

image credit: wikimedia commons

Crop wild relatives are a hot topic these days. That’s because feeding a growing population in an increasingly globalized world with the threat of climate change looming requires creative strategies. Utilizing wild relatives of crops in breeding programs is a potential way to improve yields and address issues like pests and diseases, drought, and climate change. While this isn’t necessarily a new strategy, it is increasingly important as the loss of biodiversity around the globe threatens many crop wild relatives. Securing them now is imperative.

There are about 100 species in the genus Lactuca. Most of them are found in Asia and Africa, with the greatest diversity distributed across Southwest Asia and the Mediterranean Basin. The genus consists of annual, biennial, and perennial species, a few of which are shrubs or vines. Prickly lettuce (L. serriola), willowleaf lettuce (L. saligna), and bitter lettuce (L. virosa) are weedy species with a wide distribution outside of their native range. Prickly lettuce is particularly common in North America, occurring in the diverse habitats of urban areas, natural areas, and agricultural fields. It is also the species considered to be the main ancestor of today’s cultivated lettuce.

In a paper published in European Journal of Plant Pathology in 2014. Lebeda et al. discuss using wild relatives in lettuce breeding and list some of the known cultivars derived from crosses with wild species. They write that in the last thirty years, “significant progress has been made in germplasm enhancement and the introduction of novel traits in cultivated lettuce.” Traditionally, Lactuca serriola has been the primary source for novel traits, but breeders are increasingly looking to other species of wild lettuce.

bitter lettuce (Lactuca virosa) – image credit: wikimedia commons

Resistance to disease is one of the main aims of lettuce breeders. Resistance genes can be found among populations of cultivated lettuce, but as “extensive screening” for such genes leads to “diminishing returns in terms of new resistance,” breeders look to wild lettuce species as “sources of new beneficial alleles.” The problem is that there are large gaps in our knowledge when it comes to wild lettuce species and their interactions with pests and pathogens. Finding the genes we are looking for will require “screening large collections of well defined wild Lactuca germplasm.” But first we must develop such collections.

In a separate paper (published in Euphytica in 2009), Lebeda et al. discuss just how large the gaps in our understanding of the genus Lactuca are. Beginning with our present collections they found “serious taxonomic discrepancies” as well as significant redundancy and unnecessary duplicates in and among gene banks. They also pointed out that “over 90% of wild collections are represented by only three species” [the three weedy species named above], and they urged gene banks to “rapidly [acquire] lettuce progenitors and wild relatives from the probable center of origin of lettuce and from those areas with the highest genetic diversity of Lactuca species” as their potential for improving cultivated lettuce is too important to neglect.

Lactuca is a highly variable genus; species can differ substantially in their growth and phenology from individual to individual. Lebeda et al. write, “developmental stages of plants, as influenced through selective processes under the eco-geographic conditions where they evolved, can persist when plants are cultivated under common environmental conditions and may be fixed genetically.” For this reason it is important to collect numerous individuals of each species from across their entire range in order to obtain the broadest possible suite of traits to select from.

One such trait is root development and the related ability to access water and nutrients and tolerate drought. Through selection, cultivated lettuce has become a very shallow-rooted plant, reliant on regular irrigation and fertilizer applications. In an issue of Theoretical and Applied Genetics published in 2000, Johnson et al. demonstrate the potential that Lactuca serriola, with its deep taproot and ability to tolerate drought, has for developing lettuce cultivars that are more drought tolerant and more efficient at using soil nutrients.

willowleaf lettuce (Lactuca saligna) – image credit: wikimedia commons

Clearly we have long way to go in developing improved lettuce cultivars using wild relatives, but the potential is there. As Lebeda et al. write in the European Journal of Plant Pathology, “Lettuce is one of the main horticultural crops where a strategy of wild related germplasm exploitation and utilization in breeding programs is most commonly used with very high practical impact.”

Coming Up in Part Two: Can cultivated lettuce cross with wild lettuce to create super weeds?

What Is a Water Chestnut?

This question came up on a recent episode of Every Little Thing, and while I have eaten water chestnuts on numerous occasions, I realized that I have never really considered what they were or where they came from. Thanks to the folks at ELT, I am better informed. So, why not spread the wealth?

Chinese water chestnut (not to be confused with Trapa natans, which is also commonly known as water chestnut) is in the family Cyperaceae – the sedge family. Known botanically as Eleocharis dulcis, it is a member of a sizable genus collectively referred to as the spikerushes or spikesedges. Its distribution is quite expansive, spanning sections of Australia, tropical Africa, several countries in Asia, as well as islands in the Pacific and Indian oceans. It is commonly cultivated in regions outside of its native range, including in North America as a novelty crop.

Eleocharis dulcis is a perennial, aquatic plant that grows in marshes, bogs, and the margins of other wetland and riparian areas in tropical and subtropical climates. Individual plants are clumps of tall, stiff, upright, leafless stems that can grow to over one meter tall. An infloresence is borne at the tops of stems and is a short, cylindrical cluster of small, yellow-brown florets. Clumps of stems are connected via rhizomes, and in this manner dense colonies can be formed. Rhizomes also terminate in corms, which are the edible portion of E. dulcis and the part of the plant that we refer to as water chestnuts.

Chinese water chestnut (Eleocharis dulcis) growing in a bog garden – photo credit: flickr/techieoldfox

Corms are underground storage organs. They are the bases of stems that have become thick and swollen with starch. They are often covered in papery scales – which are the remnants of leaves – that help protect the corm from being damaged or drying out. Buds on the top of the corm form shoots; adventitious roots form on the bottom of the corm. Tubers, which are also modified stems and underground storage organs, differ from corms in that they have growing points at various locations along their surface rather than a single growing point at the top.

Common misconceptions are that water chestnuts are nuts or roots. They are neither. They are corms, or in other words, they are modified stem bases. Apart from that, they are vegetables. Curiously, they are vegetables from a plant family that does not produce much in the way of food for humans. Consider that the next time you eat them. You are eating a sedge.

Corm of Chinese water chestnut (Eleocharis dulcis), the edible portion of the plant – photo credit: flickr/sclereid0309

Chinese water chestnuts can be prepared in many ways, both raw and cooked. I have only had them in stir fries, but they can also be used in salads and soups or ground into flour to make water chestnut cakes. Interestingly, even when they are cooked they remain crisp. This has something to due with the special properties of their cell walls.

As an agricultural crop they are often grown in paddies in rotation with rice. With a few preparations they can also be grown at home alongside your other vegetables. Further information and instruction can be found at various locations online including Permaculture Research Institute, Missouri Botanical Garden, and Plants for a Future.

Having only eaten water chestnuts from a can, I am anxious to try fresh, raw water chestnuts. Apparently they are available at certain Asian markets. When I get my hands on some, I will let you know what I think. Follow me on Twitter or Facebook for further updates.


What are your favorite ways to eat Chinese water chestnuts? Let us know in the comment section below.

White Rot and the Quarantine Zone, revisited

This is a revised version of a post I wrote in July 2013 during the inaugural year of Awkward Botany.


It’s garlic planting season in the northern hemisphere. A few years ago, while helping out with the garlic harvest at a local farm, I had the chance to learn about some of the challenges involved in growing garlic in southern Idaho. Apart from the fact that it is a very labor intensive crop to grow, one of the major challenges stems from a disease called white rot – easily one of the worst diseases that garlic and onion growers face.

White rot is caused by a fungus (Sclerotium cepivorum), and it affects all plants in the Allium genus, including garlic, onions, chives, and ornamentals. The disease causes the leaves of alliums to die back, their bulbs to decay, and their roots to rot, ultimately turning the plants to mush. Sclerotia, the dormant stage of the fungus, are small (about the size of a poppy seed), black, spherical structures that can survive in soil for more than 20 years. They remain dormant until the exudates of allium plants awaken them, at which point they begin to grow, unleashing their destruction. Sclerotia can be moved around by farm equipment, floods, irrigation water, wind, and by attaching themselves to plant material. Once this fungus has established itself in a field, it is extremely difficult to eradicate, making the field virtually unfit for allium crops.

The threat of white rot and the monetary damage that it can cause led to the establishment of a quarantine zone in southern Idaho in order to protect its $55 million dollar a year onion industry. Due to the quarantine zone (which encompasses 21 counties), all garlic that is grown for seed within the zone must be inspected and certified. [“Seed” in this case refers to the garlic cloves themselves; onions, on the other hand, are grown from actual seeds and are not subject to the same protocol.] Any seed garlic that is brought into the zone must go through a rigorous testing process in order to ensure that it is free of the white rot pathogen before it can be planted. Garlic is a specific threat because the cloves can readily carry sclerotia, compared to onion seeds, which are not likely to harbor them.

This process significantly limits the amount and variety of garlic that can be grown in the quarantine zone. While the quarantine is essential for warding off the threat of this particular pathogen, it stifles the garlic growing industry and makes it difficult for new garlic growers to establish themselves.

Garlic farming is already incredibly demanding due to the amount of time and physical labor that goes into planting, harvesting, drying, grading, etc. The quarantine, while understandable, is an added challenge. Learn more about this issue by listening to this story on PRX.

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Garlic emerging in the spring.

When Sunflowers Follow the Sun

Tropisms are widely studied biological phenomena that involve the growth of an organism in response to environmental stimuli. Phototropism is the growth and development of plants in response to light. Heliotropism, a specific form of phototropism, describes growth in response to the sun. Discussions of heliotropism frequently include sunflowers and their ability to “track the sun.” This conjures up images of a field of sunflowers in full bloom following the sun across the sky. However cool this might sound, it simply doesn’t happen. Young sunflowers, before they bloom, track the sun. At maturity and in bloom, the plants hold still.

What is happening in these plants is still pretty cool though, and a report published in an August 2016 issue of Science sheds some light on the heliotropic movements of young sunflowers. They begin the morning facing east. As the sun progresses across the sky, the plants follow, ending the evening facing west. Over night, they reorient themselves to face east again. As they reach maturity, this movement slows, and most of the flowers bloom facing east. Over a series of experiments, researchers were able to determine the cellular and genetic mechanisms involved in this spectacular instance of solar tracking.

Helianthus annuus (common sunflower) is a native of North America, sharing this distinction with dozens of other members of this recognizable genus. It is commonly cultivated for its edible seeds (and the oil produced from them) as well as for its ornamental value. It is a highly variable species and hybridizes readily. Wild populations often cross with cultivated ones, and in many instances the common sunflower is considered a pesky weed. Whether crop, wildflower, or weed, its phototropic movements are easy to detect, making it an excellent subject of study.

Researchers began by tying plants to stakes so that they couldn’t move. Other plants were grown in pots and turned to face west in the morning. The growth of these plants was significantly stunted compared to plants that were not manipulated in these ways, suggesting that solar tracking promotes growth.

The researchers wondered if a circadian system was involved in the movements, and so they took sunflowers that had been growing in pots in a field and placed them indoors beneath a fixed overhead light source. For several days, the plants continued their east to west and back again movements. Over time, the movements became less detectable. This and other experiments led the researchers to conclude that a “circadian clock guides solar tracking in sunflowers.”

Another series of experiments helped the researchers determine what was happening at a cellular level that was causing the eastern side of the stem to grow during the day and the western side to grow during the night. Gene expression and growth hormone levels differed on either side of the stem depending on what time of day it was. In an online article published by University of California Berkeley, Andy Fell summarizes the findings: “[T]here appear to be two growth mechanisms at work in the sunflower stem. The first sets a basic rate of growth for the plant, based on available light. The second, controlled by the circadian clock and influenced by the direction of light, causes the stem to grow more on one side than another, and therefore sway east to west during the day.”

The researchers observed that as the plants reach maturity, they move towards the west less and less. This results in most of the flowers opening in an eastward facing direction. This led them to ask if this behavior offers any sort of ecological advantage. Because flowers are warmer when they are facing the sun, they wondered if they might see an increase in pollinator visits during morning hours on flowers facing east versus those facing west. Indeed, they did: “pollinators visited east-facing heads fivefold more often than west-facing heads.” When west-facing flowers where warmed with a heater in the morning, they received more pollinator visits than west-facing flowers that were not artificially warmed, “albeit [still] fewer than east-facing flowers.” However, increased pollinator visits may be only part of the story, so further investigations are necessary.


I’m writing a book about weeds, and you can help. For more information, check out my Weeds Poll.

Summer of Weeds: Lambsquarters

Since we seem to be on the topic of edible weeds we may as well discuss lambsquarters, another frequently present and commonly eaten, nutritious and versitile weed. Botanically known as Chenopodium album, it is a member of the family Amaranthaceae and therefore related to several common (and uncommon) agricultural crops, including spinach (Spinacia oleracea), beets (Beta vulgaris), Swiss chard (also Beta vulgaris), amaranth (Amaranthus spp.), and red orach (Atriplex hortensis). Chenopodium, a genus consisting of 100 plus species, is also cultivated in various parts of the world for its edible leaves, stems, and seeds. Chenopodium quinoa, commonly known as quinoa, is now a popular “grain” in North America after being grown for millenia by Andean cultures.

Chenopodium album is a summer annual that reaches up to 6 feet tall with sturdy, angular stems and triangular, diamond-shaped, or lance-shaped leaves with irregularly toothed margins. The leaves are green on top and mealy gray-white on bottom. The flowers are tiny, petal-less, and organized in tight clusters at the ends of branches. In Botany In a Day, Thomas Elpel describes the flowers as “little green ‘globs’ forming along an upright stalk, sometimes colored with specks of yellow.” They are generally wind-pollinated, but are occassionally visited by pollinating insects. Each plant can produce tens of thousands of seeds, which are potentially viable for up to 40 years.

Inflorescence of lambsquarters (Chenopodium album)

Lambsquartes is one of many common names for C. album (others include goosefoot, fat hen, baconweed, mealweed, frostblite, and wild spinach), and is a name with several proposed origins. Is it because the plant is commonly found growing in the manure-rich soils of barnyards? Or is it because the fuzzy undersides of the leaves are reminiscent of sheep’s wool? Perhaps it is because per weight, the harvested plants and a quarter of lamb contain roughly the same amount of protein? Who knows? Despite all this talk of sheep, however, large quantities of lambsquarters are reported to be poisonous to both sheep and pigs.

Though lambsquarters prefers nutrient-rich soils, it tolerates a wide variety of soil types, including dry, compacted, urban soil. It is drawn to all sorts of disturbed sites and is particularly abundant in gardens, agricultuaral fields, and roadsides. It readily hybridizes with other Chenopodium species, including the North American native C. berlandieri. In The Book of Field and Roadside, John Eastman calls it “one of the wold’s most abundant and noxious weeds,” because “it competes with some 40 crops [and] is especially invasive in tomato, potato, sugar beet, soybean, and corn fields.”

Eastman goes on to hint at lambsquarters’ potential for phytoremediation: “The plant accumulates high levels of nitrates and pesticides in addition to its oxalic acid content.” It also takes up heavy metals, including zinc, copper, and lead. This phenomenon is worth a future post, so stay tuned.

Leaf of lambsquarters (Chenopodium album)

That being said, when harvested from a non-polluted site, lambsquarters is a very nutritious spinach-like green both raw and cooked. Younger leaves and plants are preferred because older ones tend to be higher in oxalic acid. The seeds are also edible and, like quinoa, can be used in a similar manner as common grain and cereal crops. Harvester ants and various bird species also collect and consume the seeds. The roots of lambsquarters are high in saponin and can be used to make soap.

There are many reasons to be impressed with Chenopodium album, including its ability to tolerate drougt and frost, its adaptability to all types of soil, its highly nutritious plant parts (but also potentially toxic due to accumalation of pollutants and oxalic acid), and its competitive and persistent nature. Ehrenfried Pfeiffer, author of Weeds and What They Tell, was in awe of this “most enduring annual weed” and its goosefoot family relatives, writing: “We have the feeling that the goosefoot was destined to play a better role than to become an obnoxious weed. They follow closely man’s steps, showing their inclination to be domesticated. Probably future plant breeders may develop new cultivated varieties out of this family long after our present cultivated plants have degenerated, for it is their extreme vitality and preserverence to grow that makes the goosefoot family so interesting.”

Pfeiffer’s predictions haven’t quite come to pass, but time will tell.

More lambsquarters flowers


According to an article posted on LiveScience, lambsquarters is one of “The Five Healthiest Backyard Weeds.” The list includes two other weeds we have covered during the Summer of Weeds: Purslane and Plantain.

Summer of Weeds: Eating Purslane

If it wasn’t so prolific and persistent, purslane would probably be a welcome guest in our vegetable gardens and edible landscapes. Easily among the most nutritious and versatile of the edible weeds, Portulaca oleracea is an annoyingly abundant annual that has inserted itself into garden beds and croplands in temperate climates across the globe. Thought to have originated in India or somewhere in Eurasia, purslane invaded North America long before Europeans did and has been naturalized across much of the continent for hundreds of years.

common purslane (Portulaca oleracea)

There are over 100 known species in the genus Portulaca, the only genus in the family Portulacaceae (otherwise known as the purslane family). Common purslane is a succulent plant with paddle- or teardrop-shaped leaves that generally grows low to the ground, forming a thick mat. It reaches for the sky when grown in shade or when competing with other plants for space. It produces little, yellow flowers that only open in bright sun and are typically self-pollinated. A small capsule containing dozens of tiny, black seeds quickly follows each flower. Each plant can produce tens of thousands of seeds, which remain viable for around 40 years.

Attempts to remove purslane by cultivation may only aid its survival. Broken pieces of the plant can take root in the soil, and uprooted plants can re-root if they are in contact with soil. Stirring up the ground brings to the surface seeds from purslane’s extensive seed bank. These freshly exposed seeds can then germinate, taking advantage of disturbance and open space. For all these reasons and more, John Eastman writes in The Book of Field and Roadside: “Purslane knows how to live and linger.”

The ever-urban and ever-common purslane.

The seeds of purslane germinate in late spring and throughout the summer when the soil has reached at least 75 – 80° Fahrenheit. It is adapted to high heat and dry soils. In order to conserve water, it switches to CAM photosynthesis when conditions are particularly hot and dry. In this photosynthetic pathway, carbon dioxide is stored as malic acid during the night and then converted back during the day. This means that, when it comes to eating purslane, the flavor changes depending on when the plant is harvested. In The Wild Wisdom of Weeds, Katrina Blair discusses this phenomenon: “In the morning purslane leaves contain as much as ten times more malic acid, making them very sour tasting. If you prefer a milder tasting purslane, harvest your greens in late afternoon and if you want more zing to your recipes, gather the leaves at dawn.”

Speaking of eating purslane, if all the claims are to be trusted, there may not be a more nutritious weed. In A Feast of Weeds, Luigi Ballerina calls it “a health bomb” because “it contains more omega-3 fatty acids than almost any other green, not to mention vitamins A, B, and C and beta carotene.” Blair calls it “one of the most nutritious plants on Earth,” and goes on to sing praises about its richness in dietary fiber, vitamins, minerals, protein, etc. Funnily enough, in describing the health benefits of purslane, Ballerina also quotes ancient sources claiming that “purslane calms sexual excitement.” Apparently it not only “eliminate[s] sensual dreams, but if used too much, it often extinguishes all ardor and even the capacity to procreate.”

With that caveat in mind, I tried it anyway. I had eaten it before, but nothing more than a leaf here and there and once in a green salad. I picked two recipes to try: Walnut Purslane Coleslaw from The Wild Wisdom of Weeds and Potatoes and Purslane from A Feast of Weeds. I’m generally a big fan of coleslaw, but for whatever reason I found this recipe to be a little bland. It was missing something, but I couldn’t put my finger on it. The purslane seemed to add a vague slimy-ness to it, which it will do on account of its mucilaginous nature.

Walnut Purslane Coleslaw

The Potatoes and Purslane recipe involved cooking the purslane. I enjoyed the finished product both hot and cold. The purslane added a sort of lemon-y spinach flavor. Those who tried it with me also liked it. The potato recipe was made with purslane that had been harvested in the morning, which may explain the strong lemon-y flavor. The coleslaw was made with purslane harvested in late afternoon, which may explain its blandness. I will have to try it the other way around for comparison. Purslane recipes abound in books and on the internet; browsing through them, I am intrigued enough to consider trying others. I think I’ll start with pickled purslane, purslane pesto, and perhaps, purslane sauerkraut.

Potatoes and Purslane

More Resources:


Do you have a favorite purslane recipe? Share it in the comment section below.