Article on potato leafed tomatoes and exserted stigmas

So I just came across more info on potato leaves. It’s intermingled with exserted stigma info, both of which are of interest to us folk so I’ll include that too:


Exploring Tomato Flower Structure
April 18, 2016 Seed Savers


Potato-type leaf

The world is full of gardening lore, some wise, some preposterous. Diane Ott Whealy, co-founder of Seed Savers Exchange, grew up knowing that you “harvest horseradish only in months with an ‘r’ in them,” and that “every day gets a ‘rooster step’ longer after the shortest day of the year” (Gathering 2011). The Farm Journal of 1885 suggests putting your boot alongside a melon and thumping them with your finger. If they sound the same, the melon is ripe. One common belief among gardeners is that tomato varieties with potato-type leaves are much more likely to cross pollinate than regular leaf tomatoes. What is the basis of this wisdom? Is there any truth in it? I talked to tomato breeders Tom Wagner and Craig LeHoullier to find out.

To understand the origin of this belief, we must delve into the history of tomatoes. Charles M. Rick, known as the father of tomato breeding, studied and collected seeds of wild relatives of tomatoes in their native habitat of western South America. Rick observed many differences between wild and domesticated tomatoes. Besides obvious differences such as fruit size and growth habit, Rick noted subtle but important differences in flower structure. He recognized that in wild tomatoes, the female style (the slender tube that connects the stigma and the ovary) was very long, causing the stigma to protrude much farther from the flower cone when compared to domesticated varieties (Figure 1).

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Regular leaf


Figure 1: The flower on the left has a protruding stigma (also called exserted) which sticks out past the flower cone because of the length of the flower style. The flower on the right has an inserted stigma, which is enclosed by the flower cone.

The domestication of tomatoes started in Central or South America and continued with breeding efforts in Europe and the United States, resulting in a gradual shortening of tomato flower styles. In wild tomatoes, a long style facilitated cross pollination by native insects. Outside of their native region, tomatoes had far fewer insect pollinators, so their ability to self-pollinate was essential to produce fruit. Fully inserted stigmas, which are enclosed by the flower cone, were inadvertently selected during breeding of mechanically harvested tomatoes because it ensured a reliable, concentrated fruit set (Cole 2007). Another factor in decreasing style length was breeding for heat tolerance. A study conducted by Charles M. Rick and Wesley H. Dempsey, from UC Davis, found that varieties with inserted stigmas were more heat tolerant above 35°C. All of these forces worked together to create modern varieties with short styles that were far less likely to cross pollinate than older strains that had long styles.

At the same time as tomato breeding was shortening the length of the flower style, another genetic shift was taking place: the loss of the potato-leaf trait among varieties bred for commercial production. How did this happen? Neelima Sinha of UC Davis believes that, “potato leaf was just not a trait that was important to the commercial industry so it was probably just left behind” (phone call). Given that the potato-type leaf trait is genetically recessive to regular leaves and it was much less common than regular leaves to begin with, this theory seems plausible.

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Figure 2: Normal tomato flower

However, legendary tomato breeder Tom Wagner suggests a number of hypotheses for why the loss of the potato-type leaf may have been deliberate. According to Wagner, the potato-type leaf was an oddity and did not fit the common notion of what a tomato plant looked like. Additionally, potato-type leaves may have been detrimental to plant health. Wagner says that potato-leaf plants are more prone to wind damage, fungal diseases, and uneven ripening, and the leaves obscure the fruit more than regular leaves. Today, the potato-type leaf trait represents an older class of tomatoes that was not subject to the forces working to reduce the length of the style.

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Figure 3: Marigold-type tomato flower

Another reason that potato-type leaf varieties typically have exserted stigmas has to do with genetics. Potato-leaf varieties are relatively rare and comprise a narrow genetic pool with fruit that are typically large and convoluted. These so-called beefsteak tomatoes are the result of double-flowers, sometimes called fasciated or “marigold”-type flowers (Figure 3). By nature, these flowers have more exposed, exerted stigmas. Hence, the potato leaf trait has become a convenient way to classify varieties that are more prone to out-crossing.

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At Seed Savers Exchange, we have been collecting evaluation data on our tomato varieties for the last several years, including data on leaf type and style length. Of the 23 potato-leaf varieties that we have recently evaluated, 16 had exserted stigmas; 6 had stigmas level with the anther cone; and one had an inserted stigma (Figure 4). This supports the belief that the potato-leaf trait is correlated with flower structure, but shows that there are exceptions to this rule. Conversely, among the varieties with regular leaves, the stigma positions were inserted, exserted, and at the same level of the anther cone in roughly equal proportions, suggesting that no conclusion can be drawn about flower structure from this leaf type (Figure 5)

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Although the potato-leaf trait may be a useful and convenient way to predict flower structure, there are exceptions. Among the potato-leaf varieties we evaluated, ‘Yellow Cherry’ had potato leaves and an inserted stigma, while several others had stigmas level with the anther cone. Another exception to the rule, according to Tom Wagner, is ‘Bloody Butcher,’ a potato-leaf type which produces red globe tomatoes and has inserted stigmas. Craig LeHoullier, tomato breeder and author, believes that “the age of the strain and the size of the fruit” are better predictors of flower structure than leaf type (e-mail correspondence).

Ultimately, the best way to determine flower structure is to actually look at the flowers themselves. It is important to note that flower structure is just one of many factors in predicting crossing rate and deciding how much to isolate a variety. Other important factors to consider are natural barriers between varieties, such as woodlands, and, especially, the number of insect pollinators that visit the flowers. Here at Heritage Farm, where varietal purity is of utmost importance, we have found that an isolation distance of 1000ft between tomato varieties almost always prevents crossing. However, Seed Savers Exchange’s collection has many more old, heirloom varieties than modern varieties and the farm has a healthy insect population with areas of woodlands and varied terrain. Every seed saver must assess these factors in their own garden or farm when deciding how much to isolate their varieties. Potato-type leaves as an indicator of flower structure is but one useful tool to help guide this decision. A much more reliable method is look at the flowers themselves.


Source: Exploring Tomato Flower Structure — Seed Savers Exchange Blog

There’s one comment on that post, and it seems worth including here:


Though fasciated fruit types play a major genetic role, exherted or exposed stigmas can also be the result of environmental conditions and other primitive genetics reintroduced by moderning breeding.
Rough flowers or double flowers can form on just about any variety if weather conditions are right. Usually these are the result of improper floral development during primordial stages (baby flower buds barely visible to the eye) in cold weather.
Anther cone (the part that envloses the stigma) development may also be damaged by insects thus exposing the style/stigma. The damage can be to the anther cone itself (usually at its base) or the developing fruit which lead to abnormal development and splitting calxes.
Styles (the long part which the stigma sits) of any variety may grow beyond the anther cone and prone to outcrossing due to hormonal changes or any such conditions which result in a sudden spurt of growth. The most common examples of this occurring happen when there is sudden change in temperature and/or sudden uptake in water and/or fertilizer. Hormonal issues from faint amounts of herbicide drift may also in some cases result in exherting stigmas (2,4-D can drift for miles in the wind).

More modern varieties that have been crossed to genetically diverse primitive or wild types in their development (usually to get disease resistance) may also exhert as some wild types are not self-compatible. Though there is effort to breed this out, it may show up under stress.
Concerning potato-leaf types: There has been breeding work done in the past with them. I came across mention of them in a “sand damage resistance” study being done in Florida as part of the STEP project (Southern Tomato Exchange Program - whereas primarily 50-60’s in the southern part of the US, breeders freely exchanged lines between their programs for evaluation). This study referenced lines from Oklahoma bred for “wind resistance” as the leaves protected the fruits from blowing sand and dirt. I was never able to trace down the lines.
Like the USDA with their PI numbers for named varieties, the STEP program seemed to have had their own numbering system (example: the Univ of Missouri tomato named “Advanced” was noted as STEP 667. In STEP, this tood for “39-4-36” as it was called in Univ or Missouri records. One needed to know that Missouri line to cross reference that STEP 667 = “Advance”). I have searched for years about asked several breeders if they knew of a key for the STEP lines. However most of these breeders came after that programmed had ceased and did not know of any specific record of a list. It may exist somewhere in a university’s stack or files retained from those professors. Unfortunately some of that stuff gets thrown out, like at retirement (I know, I helped throw away old notebooks and data on bean genetics once).
Both of us being from Kansas, and knowing “wind”, I mentioned this study to Tom (Wagner) in conversation. Maybe he heard wrong or forgot. It does stand to reason that larger leaves would create a greater surface area for wind resistance (think of a larger sail on a boat catching more wind). Such resistance could be an issue for poorly supported plants. This issue here was protecting the fruit from damage of particles in the wind. That makes sense. My guess is that idea fell out of favor by just creating more yield in other lines and making up for such potential losses. That seems to be a prevailing thought in much of the commercialized breeding programs.
I heard repeatedly in the past that commercial growers did not like potato leafed types simply because “thats not the way they are supposed to look.” Things change though. Some old timers in the Carolina’s did NOT like uniform green shouldered fruit (the shoulders are light green vs dark in the unripe stage but ripen uniformly red). More recent Israeli research has pointed out that breeding for the uniformly green shoulders trait is one of the factors which has reduced tomato flavor. Apparently the Carolina old timers were onto something.

Also note that this part is a bit misleading:

That’s actually not always the case. Many accessions of pimpinellifolium, cheesmaniae, and galapagense, are not exserted - perhaps most? Chmielewskii and neorickii are both self compatible also, and whilst chmielewskii is said to be always exserted, I’m not sure about neorickii.

And I’m quite surprised about this part:

Is that representative of domestic tomatoes in general? I’ve very surprised one third were exserted and only about one third inserted!

interesting, potato leaf are used a lot in breeding projects (I know the dwarf breeding project relied on them a lot) because it’s a recessive trait. So a female potato leaf with a male tomato leaf if the seeds have tomato leaf then you know the cross was successful. Seems like that could help us here too, though I assume that in the long run the tomato leaf would take over in the breeding project unless it really is a requirement

More info from ‘Rates of cross-pollination in Lycopersicon pimpinellifolium: Impact of genetic variation in floral characters’ by Charles M. Rick et. al.:

Cross-pollination was tested between different genotypes of Lycopersicon pimpinellifolium in an experimental plot situated in the southerly range of the species. Rates of cross-pollination were found to be highly correlated with flower size and were related to a lesser extent with degree of stigma exsertion. Comparisons of fertility suggest that exsertion of 1.5 mm or more from the anther tube orifice interferes with automatic self-pollination, thereby increasing rates of cross-pollination.

That seems consistent with what I have been noticing. If you pick the plant with the longest exsertion you get the most cross pollination.

Sure. But what I found surprising was the 1.5mm limit (I should really pay attention to that!) and that flower size was more relevant than exsertion. Though, presumably at least some exsertion (or at least no insertion) would be necessary for flower size to make much of a difference?

Still, I’m wondering if exsertion just might be an unsustainable aim so long as there’s pressure from non-exserted neighbours. What do you reckon - that would slowly over generations breed out the exsertion, no? I mean, even if it would take 30 or 50 years - but if we’re thinking longterm like that - or actually that’s maybe what I’d call medium term!

I think that as long as I have at least one potato leaf plant with extreme exsertion each year planted very close to other regular leaf tomatoes with good pollen production I’ll be ok for at least a few new F1 hybrid plants the next year. The mother plant produces some selfed and some crossed offspring and you can let the hybrids segregate back to potato leaf the next year.

If you’re saying exsertion to be unsustainable long term due to your plants being pollinated by your neighbor’s tomatoes then Mr Lofthouse has covered the math on that and it is much less likely to have pollen from your neighbor’s patch than from your patch. And you can reselect in the case it does occur.

The other thing is that if you’re crossing your plants and creating stronger plants then your neighbors are more likely to swap to what you’re growing if they see you’re having better success than them.