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.”

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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?

Is There a Place for Weeds in Urban Ecosystems?

Highly urbanized areas have a long history of disturbance. They are a far cry from the natural areas they once displaced, bearing little resemblance to what was there before. In this sense, they are a brand new thing. During the urbanization process, virtually everything is altered – temperatures, soils, wind patterns, hydrology, carbon dioxide levels, humidity, light availability, nutrients. Add to that a changing climate and increased levels of pollution, and the hope of ever seeing such a site return to its original state – whatever that might mean – is crushed.

What then should we consider the natural flora of an ecosystem like this? Certainly it is not the native flora that once stood on the site before it was developed; virtually none of the conditions are the same anymore. If we are defining “natural” as existing with minimal human intervention, then the natural urban flora would be whatever grows wild outside of our manicured landscapes and managed, remnant natural areas. It would be a cosmopolitan mixture of plants that have joined us in our migrations with and without our permission, along with a collection of species that are either extant to the site or have been brought in by wildlife. In many ways it would mirror the human populations of our modern cities – an assortment of residents from around the globe with diverse backgrounds and cultural histories.

In Wild Urban Plants of the Northeast, Peter Del Tredici classifies urban land into three general categories based on their ecological functions: native, remnant landscapes; managed, constructed landscapes; and ruderal, adaptive landscapes. Native, remnant landscapes are generally small areas within city limits that have never been developed. They contain a portion of the native plants that once populated the area, and they require vigilant and regular maintenance to keep non-native plants from invading and to control those that already have. Managed, constructed landscapes include all of the parks and gardens that have been designed and intentionally planted. They require regular maintenance of varying intensity in order to keep them looking the way they are intended to look. Ruderal, adaptive landscapes are abandoned or neglected sites that are populated by plants that have arrived on their own and that maintain themselves with virtually no human intervention. This is where the true, wild urban flora resides.

Prickly lettuce (Lactuca serriola) growing in an abandoned lot.

Many of the plants that make up our wild urban flora are what we commonly refer to as weeds. These weedy plants appear in landscapes throughout our cities, but are generally removed or controlled in all landscapes except the abandoned ones. It is in these neglected sites that weeds have the greatest potential to provide vital ecosystem services, performing ecological functions that are beneficial to urban life.

Not all plants are suited for this role. Spontaneous urban vegetation is particularly suited due to its ability to thrive in highly modified, urban environments without external management. Regardless of provenance, this suite of plants, as Del Tredici points out, seem to be “preadapted” to urban conditions and “are among the toughest on the planet.” A long list of traits has been identified for plants in this category, ranging from seed dispersal and viability to speed of growth and reproduction to tolerance of harsh conditions. Del Tredici summarizes by stating, “a successful urban plant needs to be flexible in all aspects of its life history from seed germination through flowering and fruiting, opportunistic in its ability to take advantage of locally abundant resources that may be available for only a short time, and tolerant of the stressful growing conditions caused by an abundance of pavement and a paucity of soil.”

Abandoned lots flush with weeds, overgrown roadsides and railways, and neglected alleyways colonized by enterprising plants are generally seen as ugly, unsightly eyesores – products of neglect and decline. Some of the plants found in such locations are valued in a garden setting or prized as part of the native landscape in a natural area, but growing wildly among trash and decaying urban infrastructure they, too, are refuse. As Richard Mabey has written: “If plants sprout through garbage they become a kind of litter themselves. Vegetable trash.”

Abandoned chicken coop overtaken by tree of heaven saplings (Ailanthus altissima).

Despite how we feel about these plants or the aesthetics of the locations they find themselves in, they are performing valuable services. Apart from adding to the biodiversity on the site as well as producing oxygen and sequestering carbon – services that virtually all plants offer – they may be preventing soil erosion, stabilizing waterways, absorbing excess nutrients, reducing the urban heat island effect, mitigating pollution, building soil, and/or providing food and habitat for urban wildlife. While cultivated and managed landscapes can achieve similar things, these neglected sites are doing so without resource or labor inputs. They are sustainable in the sense that their ability to provide these services is ongoing without reliance on outside maintenance.

Sites like these should be further investigated to determine the full extent of the services that they may or may not be offering, and in the event that they are doing more good than harm, they should be conserved and encouraged. One service that is receiving more attention, as Del Tredici writes, is phytoremediation – “the ability of some plants to clean up contaminated sites by selectively absorbing and storing high concentrations of heavy metals such as cadmium, lead, copper, zinc, chromium, and nickel in their tissues.” Weed species with this ability include prickly lettuce (Lactuca serriola), lambsquarters (Chenopodium album), and mugwort (Artemisia vulgaris). In an article in Places Journal, Del Tredici gives the example of the often despised, introduced plant, common reed (Phragmites australis) cleaning up the New Jersey Meadowlands by “absorbing abundant excess nitrogen and phosphorous throughout this highly contaminated site.”

In the book, Weeds: In Defense of Nature’s Most Unloved Plants, Richard Mabey writes: “As we survey our long love-hate relationship with [weeds], it may be revealing to ponder where weeds belong in the ecological scheme of things. They seem, even from the most cursory of looks, to have evolved to grow in unsettled earth and damaged landscapes, and that may be a less malign role than we give them credit for.” Perhaps, seeing them in this worthy role, will temper our knee-jerk inclination to demonize them at every turn.

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See Also: Our Urban Planet and Wild Urban Plants of Boise.