Domestication influences interactions between mycorrhizae and solanaceous plants in agricultural settings

Domestication influences interactions between mycorrhizae and solanaceous plants in agricultural settings | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 9 March 2026 V1 Latest version Share on Domestication influences interactions between mycorrhizae and solanaceous plants in agricultural settings Authors : Zoe Getman-Pickering 0000-0002-5695-858X [email protected] , Jennifer Thaler , Jules Davis 0000-0002-4902-675X , Heather Grab , Sheyla Finkner , Elyse McCormick , Abigail Getman-Pickering , and Kaitlin Deutsch Authors Info & Affiliations https://doi.org/10.22541/au.177307596.64983135/v1 141 views 63 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract During domestication, selection has typically decreased plant resistance to herbivores and reduced beneficial relationships with microbial mutualists such as mycorrhizae. These two processes raise the question of whether reductions in plant-mycorrhizal symbioses during domestication altered mycorrhizae-conferred resistance to herbivores. We conducted a factorial experiment using 6 pairs of domesticated and wild solanaceous crop species, with and without mycorrhizae. We measured growth, physical and chemical resistance traits, and resistance to three species of herbivorous insects. Mycorrhizae strongly affected undomesticated plants but had no effect on domesticated plants. In undomesticated plants, mycorrhizae reduced plant growth both above and below ground, altered resistance to generalist herbivores (Podisus maculiventris, Manduca sexta, and Trichoplusia ni), and increased plant protease inhibitors, a defensive metabolite. However, these effects were completely absent for domesticated plants. The lack of effect of this mycorrhizal inoculum in domesticated plants suggests changes in expectations of broader plant-mycorrhizae mutualisms in domesticated plants and calls into question the value of mycorrhizae as an agricultural tool. The disrupting of the plant-mycorrhizal symbiosis may be a new mechanism by which domestication has decreased plant defenses and resistance to herbivores. Abstract During domestication, selection has typically decreased plant resistance to herbivores and reduced beneficial relationships with microbial mutualists such as mycorrhizae. These two processes raise the question of whether reductions in plant-mycorrhizal symbioses during domestication altered mycorrhizae-conferred resistance to herbivores.We conducted a factorial experiment using 6 pairs of domesticated and wild solanaceous crop species, with and without mycorrhizae. We measured growth, physical and chemical resistance traits, and resistance to three species of herbivorous insects.Mycorrhizae strongly affected undomesticated plants but had no effect on domesticated plants. In undomesticated plants, mycorrhizae reduced plant growth both above and below ground, altered resistance to generalist herbivores ( Podisus maculiventris , Manduca sexta, and Trichoplusia ni ), and increased plant protease inhibitors, a defensive metabolite. However, these effects were completely absent for domesticated plants.The lack of effect of this mycorrhizal inoculum in domesticated plants suggests changes in expectations of broader plant-mycorrhizae mutualisms in domesticated plants and calls into question the value of mycorrhizae as an agricultural tool. The disrupting of the plant-mycorrhizal symbiosis may be a new mechanism by which domestication has decreased plant defenses and resistance to herbivores. Word count: 5652 words Key words: Domestication, mycorrhizae, herbivory, chemical ecology, plant-herbivore interactions, tri-trophic interactions, above-belowground interactions Introduction During the process of crop domestication, plants face novel changes in selective forces (Meyer et al. , 2012; Turcotte et al. , 2017) Human-mediated selection typically results in increased plant growth (Pickersgill, 2007) and nutritional content, decreased defensive chemistry (Lindig-Cisneros, 1997; Whitehead et al. , 2017; Luna-Ruiz et al. , 2018), and altered interactions with mutualists (Turcotte et al. , 2014; Pérez-Jaramillo et al. , 2016 and citations therein) and herbivores (Chen et al. , 2015; Whitehead et al. , 2017 and citations therein). Domesticated plants tend to be more susceptible to herbivores (Whitehead et al., 2017) which has been attributed to changes in different selective forces including selection for increased growth, selection for increased nutrient quality, and selection for more palatable plants. First, selection for faster growing and larger plants might lead to reduced resistance to herbivores as plants funnel resources away from defense and into producing edible tissue (Huot et al., 2014), assuming a growth-defense tradeoff (Rosenthal & Dirzo, 1997). Secondly, humans have selected for more nutritious plants, which may make the plants more attractive to herbivores (Delgado-Baquerizo et al. , 2016). Finally, humans have selected for lower levels of bitter and unpalatable defense compounds, making the plants more palatable for both humans and insects (Benrey et al. , 1998; Whitehead et al. , 2017; Chomicki et al. , 2020). Beyond that, human pest control efforts can alleviate selective pressure by herbivores, further reducing the need for defense (Macfadyen & Bohan, 2010). In sum, these effects result in lower defenses and lower resistance to herbivores in domesticated plants compared to their wild relatives (Whitehead et al. , 2017). In addition to changes in growth and resistance, the domestication process has altered many crop plants’ relationships to microbial symbionts (Mutch & Young, 2004; Zachow et al. , 2014; Coleman-Derr et al. , 2016). A ubiquitous microbial symbiont, mycorrhizae are endophytic fungi that provide nutrients in exchange for photosynthate, increasing their host plant’s growth and survival. As seen with symbionts more broadly, domesticated plants tend to form less robust associations with mycorrhizae. In a manipulative greenhouse study Turrini et al., (2016) found that domesticated sunflower ( Helianthus annuus ) plants had lower instances of mycorrhizal colonization than their wild counterparts. However, the results were variable, and it was not tested whether the sunflower plants benefited from the association. In a field survey, assessment of over 50 breadfruit ( Artocarpus altilis ) cultivars found that again, domesticated plants had lower colonization and lower diversity of colonizing fungi compared to wild and landrace varieties (Xing et al. , 2012). An impressive and comprehensive greenhouse study of 27 domesticated crop species and their wild progenitors found that the wild progenitors had higher mycorrhizal colonization and growth than their domesticated counterparts unless fertilizer levels were very low (Martín‐Robles et al. , 2018). While reduced association with mycorrhizae appears common, it is unclear how the reduction in symbioses in domesticated plants might affect mycorrhizae-conferred resistance to insects. Besides altering plant growth, mycorrhizae can also alter plant resistance traits and thus resistance and susceptibility to herbivores. However, the effects are highly variable depending on abiotic conditions and the identities of the organisms involved (Gange & West, 1994; Pozo et al. , 2002; Koricheva et al. , 2009). There is ample evidence in a wide range of systems of both mycorrhizae-conferred resistance (Gange & Nice, 1997; Gange, 2001; Wooley & Paine, 2007), and mycorrhizae-conferred susceptibility to herbivores and diseases (Gange et al. , 1999, 2005; Babikova et al. , 2013). Mycorrhizae frequently confer resistance to generalist herbivores and mesophyll-feeding, piercing-sucking insects, while mycorrhizae confer susceptibility to specialist herbivores and phloem feeders (Koricheva et al. , 2009). Furthermore, mycorrhizae alter resistance and mechanisms of resistance differently in different plant species. A study of two undomesticated Solanum species found that mycorrhizae altered defenses against herbivores in only one of the two species (Minton et al. , 2016). It is unclear whether related domesticated and non-domesticated plants show similar changes to susceptibility or resistance to herbivores when colonized by mycorrhizae. Mycorrhizae alter their host plant’s resistance to herbivores through mechanisms including altering expression of defensive compounds, changing host plant nutrient quality, and by making plants more or less attractive to natural enemies. A plant can respond to mycorrhizal colonization by inducing jasmonate and salicylate signaling pathways (Pozo & Azcón-Aguilar, 2007; Jung et al. , 2012) and thus upregulating the expression of plant defenses controlled by those pathways, such as trypsin protease inhibitors (Song et al. , 2013, 2014, Getman-Pickering et al. 2020) and trichomes (Malik et al. , 2018). By increasing the absorption of macro- and micronutrients, mycorrhizae can increase nitrogen and decrease the carbon:nitrogen ratio in their host, making the plant more attractive to herbivores. However, as domesticated plants tend to lower association with mycorrhizae, we might predict that mycorrhizae will have less effect on resistance traits and resistance to herbivores in domesticated plants. Domestication and association with mycorrhizae can each independently change plant resistance to herbivores. We predicted that domestication has reduced the relationship between mycorrhizae and their host plants and has thus reduced mycorrhizae-conferred resistance to herbivores in these domesticated plants. Our key goal in this study was to evaluate the impact of a mycorrhizal inoculum on resistance and biomassin 6 pairs of domesticated and undomesticated solanaceous plants. Specifically, we asked the following questions: 1) Has domestication altered the degree to which crop plants are colonized by mycorrhizae? 2) Has domestication altered the degree to which plant growth benefits from mycorrhizae? 3) Do domestication and mycorrhizae interact to alter the host plant’s physical and chemical resistance traits? 4) How has domestication altered mycorrhizae-conferred resistance to herbivores in the host plants? Design To test whether domestication has changed mycorrhizae-conferred resistance to herbivores, we used a paired approach with domesticated and undomesticated progenitors of 6 common solanaceous crops. This paired crop design (similar to Martin-Robles 2018) allowed us to reduce noise from interspecific variation in our statistical analyses. Using a fully factorial design, the paired domesticated and undomesticated crops were grown with and without a single mycorrhizal inoculum (see Mycorrhizal inoculation section) . Each of these plants was grown with either low, medium, or high phosphorus fertilizer for a total of 72 treatments (6 crops x 2 domestication levels x with or without mycorrhizae x 3 fertilizer treatments) (n=11-12 plants per treatment, 835 total). Due to a lack of substantial effect, the methods and results of the fertilizer treatment have been moved to the Supplemental Information 2. This experiment was conducted in a single experimental run from July-November 2018. Plants We selected 6 agronomically important solanaceous crops: tomato, potato, eggplant, chili, tomatillo, and tobacco. For each solanaceous crop, we obtained seeds of two accessions: one representative of a domesticated genotype, and one of its recognized undomesticated progenitors. For tomatillo ( Physalis philadelphica ), potato ( Solanum tuberosum ), and chili ( Capsicum annum ) we used an undomesticated accession of the same species. For tomato ( S. lycopersicum ), eggplant ( S. melongena ) and tobacco ( Nicotiana tabacum ) we used closest known available undomesticated relatives from a different species S. pimpinellifolium , S. linnaeanum , and N. sylvestris respectively. Detailed information about seed accessions are in Appendix 1. Sterilization and Germination We grew the plants in a 1:1 sand and calcined clay media (Turface™). To ensure there were no naturally occurring mycorrhizal communities in the media, we autoclaved the media for one hour 3 times, 24 hours apart at 121 °C. To recover soil microbes, which could have an impact on both plant health (Berendsen et al., 2012 and references therein), mycorrhizal fungi (Desirò et al., 2014), and the interaction between the two, we filtered a mixture of Lambert LM-AP potting soil and water through a 1 micron sieve, and added 20 mL of the resulting solution to each pot. We used potting soil to reduce the risk of introducing pathogenic species. Seeds were surface sterilized 50:50 water bleach solution with 0.05% Triton-X. Undomesticated and domesticated tobacco seeds were surface sterilized for 4 minutes while all other seeds were surface sterilized for 30 minutes. They were then rinsed and allowed to germinate in petri dishes with moist filter paper and kept in the dark at 21 °C. After germination, the seedlings were transferred to a 10 cm pot filled with autoclaved media. Mycorrhizal inoculation Rhizophagus irregularis (previously Glomus intraradices ) is a commercially available species of mycorrhizal fungi used in agriculture and home gardens. It is highly generalist and will colonize pepper (Aissa et al., 2016, Balog et al., 2017), tomato (Formenti and Rasmann 2000, Pozo et al., 2001 Calvo-Polanco et al., 2014), tobacco (Shaul et al., 1999, Groten et al., 2015, Davis et al., 2019, Song 2019), eggplant (Douds Jr et al., 2017), tomatillo (Gómez and Margarita 2014) and potato (Hijri 2015, Alaux et al., 2018), and has been shown to alter its host’s resistance to herbivores in tomato and tobacco (Wooley and Pain 2007, Song et al 2014, Getman-Pickering 2020). A single mycorrhizal inoculum was sourced from Mycorrhizae Premier Tech P-501. After growing for 2 weeks, seedlings were randomly assigned mycorrhizae treatments (~400 plants/treatment) and inoculated with either 0.75 g of 500 spores/g inoculum or 0.75 g inoculum that was sterilized by autoclaving it for one hour 3 times, 24 hours apart at 121°C. The inoculum was suspended in water and pipetted to the base of the seedling and watered down with 50 ml of water. Harvest After transplanting, the plants were allowed to grow for 12 weeks at 27 °C in a greenhouse. All pots were kept on top of a 9cm petri dish to reduce contamination between pots and to improve water retention. All plants were kept at least 10 cm apart so they were never overshadowing each other. At the end of the experiment before harvesting, we recorded the presence and number of buds, flowers, and fruit both to monitor development time and because budding and flowering plants have different defensive strategies such as reduced induction of defenses. The remaining above and below ground biomass was harvested and dried for 3 days at 95 degrees before being weighed. Mycorrhizal Quantification To test how domestication has altered the degree to which plants associate with mycorrhizae, we measured percent colonization. 2-6 plants were randomly selected from each of our 72 treatments (~250 total) to measure mycorrhizal colonization. The roots were cleared in a 10% KOH solution and dyed using a 5% ink-vinegar solution (Sheaffer Skrip Bottled Ink, Black). For each plant, ten 1cm segments of root were randomly selected and mounted on slides for quantification with microscopy at 400x magnification using the grid intersect method developed by McGonigle et al. (1990). Defense traits Trichomes Trichomes are hairlike structures that form on the surface of leaves and contribute to plant resistance to herbivores in many plants including many solanaceous species. To determine how mycorrhizae and domestication affected trichome expression, we counted trichomes on one leaf from each plant. Due to resource constraints we only measured trichomes on ⅓ of leaves (those that received medium phosphorus fertilization, Supplemental Information) We used a dissecting microscope at 25x magnification to count the trichomes. We placed an index card with a (6+/-0.5mm) hole punched out of it on top of the leaf, avoiding the midrib and major veins. We recorded the amount of trichomes present in the 6 mm disk. Trichomes were counted on three consistent spots per leaf and the results were averaged. While trichomes of different types were measured, types varied across species, making it logistically unfeasible to perform cross species comparisons. Protease Inhibitors We evaluated the effect of domestication and mycorrhizae on plant chemical defense by measuring trypsin protease inhibitor activity. Protease inhibitors are a class of chemical defenses that reduce the digestibility of leaf tissue and can have strong negative effects on chewing herbivores (Mithöfer and Boland 2012). These defenses are both constitutive and induced (Zavala et al. 2004), Mycorrhizae have been shown to both increase and decrease protease inhibitor levels in solanaceous plants (Barazani 2005, Getman-Pickering et al. 2020 ). We excised leaves or leaflets from a consistent place on each crop type in each treatment and immediately froze the leaf tissue on dry ice. We analyzed 100 mg of tissue using a colorimetric assay to calculate the activity of defensive Trypsin Protease Inhibitors using a method adapted from Hegedus et al., (2003). Protein As many herbivores are nitrogen limited, their host choice and performance can be dependent on the availability of nitrogen and protein in their host plant’s tissue (Mattson 1980, Felton 1996). We excised leaves or leaflets from a consistent place on each crop type from every plant and immediately froze it on dry ice. We analyzed the total leaf protein using one leaf for each plant (mg/g) with a modified version of the ThermoFisher Pierce™ BCA Protein Colorimetric Assay following the manufacturer’s instructions . Herbivore assays T. ni We used the second-instar cabbage looper Trichoplusia ni (hereafter T. ni) caterpillars in a bioassay to measure plant resistance. T. ni are Noctuid caterpillars that feed on a wide variety of plant species including solanaceous and cruciferous crops (Shorey et al. 1962). These caterpillars were chosen because they are generalists that can feed on the range of plants used in this study, and because they are sensitive to changes in host plant quality. We obtained T. ni eggs from a colony maintained on artificial diet at Cornell University. The eggs were hatched, and the larvae were allowed to feed on artificial diet (Southland Products Inc) for three days. The second-instar larvae were weighed and placed in a petri dish on an excised leaf from one of the 6 plant species pairs on moist filter paper. We placed a single larva on one leaf from every surviving plant in the experiment (n=~790). The petri dishes were wrapped in parafilm to prevent desiccation and maintained in a growth chamber at 27°C with an 18/8 light cycle for three days. After three days, we noted survival and weighed the surviving larvae. We measured the leaf area consumed by each T. ni larvae using LeafByte (Getman-Pickering et al. , 2020). M. sexta Tobacco hornworm caterpillars, Manduca sexta (hereafter M. sexta), are specialist sphingid herbivores that feed on solanaceous plants. They are a common pest of tomato and tobacco in southern parts of the United States and have demonstrated resistance to solanaceous defenses (Wink & Theile, 2002; Pauchet et al. , 2010). The eggs were obtained from a colony maintained on artificial diet at Cornell University. After hatching, the caterpillars were immediately placed onto the excised leaf in a petri dish on moist filter paper. As with the T. ni , the petri dishes were wrapped in parafilm and maintained in a growth chamber for three days. After three days, we noted survival and weighed the surviving larvae. We measured the leaf area consumed by each M. sexta using LeafByte (Getman‐Pickering et al. , 2020). P. maculiventris Spined soldier bug Podisus maculiventris (hereafter P. maculiventris) are omnivorous stink bugs that feed on a wide variety of both plants and insects. P. maculiventris get most of their calories from consuming insects, however, they supplement their diet with phloem and mesophyll from leaves and stems, causing some damage to selected plants (Ruberson et al. 1986). This damage is typically minimal, as such they are commonly used as biocontrol agents in commercial crops. P. maculiventris spends much of its time walking and feeding on its host plants and is therefore affected by plant defenses more than many other natural enemies (Thaler et al. , 2015). Early instars in particular can be harmed and killed by plant trichomes (see Trichomes section). Previous work has found that trichomes can limit, incapacitate or kill early instar P. maculiventris nymphs, both through external contact with trichomes and consumption of leaf tissues (Lambert, 2007). However, it is unknown how mycorrhizae and domestication have altered the host plant quality, or how this potential change impacts P. maculiventris growth. We conducted a bioassay to determine the effect of mycorrhizae and domestication on the omnivorous natural enemy P. maculiventris . Due to time and funding constraints, we chose a single fertilizer level to analyze the effects of domestication and mycorrhizae on trichomes and P. maculiventris nymphs. This assay was only conducted on plants that were fertilized with medium phosphorus fertilizer. When the plants were ready to be harvested, we removed a leaf from the middle of a plant sample, placed it inside of a labeled 9 cm petri dish, on top of moist filter paper. Twelve hours before beginning a bioassay, we placed second-instar P. maculiventris nymphs in deli cups with a moist cotton ball for water, so they were ready to plant-feed the next day. After about 12 h of starving, P. maculiventris nymphs were weighed. We then used a paint brush to carefully place a single nymph on the center of each leaf. Every 2 hours for twelve hours we checked the location and recorded if they were on the top or bottom of the leaf or if they were on the petri dish. After 12 h we weighed the nymph and stored each leaf at -80°C for trichome counting. Statistics All statistics were conducted using R version 3.5.2. Linear models and generalized linear models were performed using lme4 (Bates et al. , 2015). In all models testing the effects of mycorrhizae and domestication on plant growth and resistance to herbivores we included mycorrhizae, domestication, fertilizer, and crop as interacting fixed effects. Four-way interactions were removed from all models as we did not have four-way predictions to test. Post-hoc LS means comparisons were conducted using emmeans (Lenth 2019), and pairwise comparisons of crops were corrected using FDR. We ran pre-planned pairwise comparisons of the effect of mycorrhizae within domestication treatments on most models. We did not run pairwise comparisons on Fertilizer by Crop interactions regardless of significance as these interactions were not relevant to our questions. Models, distributions and transformations can be found in Table 1 below. Additionally, when protein content was included as a predictor variable it was rescaled by dividing by 100. A full table of all models, results, and post-hoc results can be found in Supplemental Information 3. Plots were produced by ggplot2 using the model outputs (Wickham, 2016). Table 1 A list of statistical models used to analyze the data presented here. Distributions and transformations were selected to meet model assumptions. Response variable Predictor Variable Model Random Effects Distribution Response Variable Transformation Percent Colonization Domestication*Fertilizer* Crop cbind Generalized Linear Model n/a quasibinomial n/a Above Ground Biomass Domestication*Mycorrhizae*Fertilizer * Crop Linear Model n/a linear Sqrt Below Ground Biomass Domestication*Mycorrhizae*Fertilizer*Crop Linear Model n/a linear Sqrt Flowering Time Domestication*Mycorrhizae*Fertilizer*Crop GLM n/a binomial n/a Protease Inhibitors (percent inhibition) Domestication*Mycorrhizae*Fertilizer*Crop cbind GLM n/a quasibinomial n/a Protein Domestication*Mycorrhizae*Fertilizer*Crop LM n/a linear Sqrt Trichomes Domestication*Mycorrhizae*Fertilizer*Crop GLM n/a quasipoisson n/a T. ni Leaf Area Consumed Domestication*Mycorrhizae*Fertilizer*Crop LM n/a linear Sqrt T. ni Leaf Area Consumed Protein*Protease Inhibitors LM Crop linear Log T. ni Percent Weight Change Domestication*Mycorrhizae*Fertilizer*Crop LM n/a linear Log T. ni Percent Weight Change Protein*Protease Inhibitors LM Crop linear n/a T. ni Survival Domestication*Mycorrhizae*Fertilizer*Crop GLM n/a quasibinomial n/a T. ni Survival Protein*Protease Inhibitors GLMM Crop quasibinomial n/a M. sexta Leaf Area Consumed Domestication*Mycorrhizae*Fertilizer*Crop LM n/a linear Sqrt M. sexta Leaf Area Consumed Protein*Protease Inhibitors LME Crop linear n/a M. sexta Percent Weight Change Domestication*Mycorrhizae*Fertilizer*Crop LM n/a linear Sqrt M. sexta Percent Weight Change Protein*Protease Inhibitors LME Crop linear n/a M. sexta Survival Domestication*Mycorrhizae*Fertilizer*Crop GGLM n/a binomial n/a M. sexta Survival Protein*Protease Inhibitors GLMM Crop binomial n/a P. maculiventris Percent Weight Change Domestication*Mycorrhizae*Fertilizer*Crop LME n/a linear Log P. maculiventris Percent Weight Change Protein*Protease Inhibitors*Trichomes LME Crop linear Log Results Colonization The effects of domestication were dependent on crop (deviance=-207, df=107, p=0.012). Domesticated crop plants had 44-75% higher, or marginally higher colonization than undomesticated plants for all crops except Physalis (Supplemental Information 3). Plant growth traits Mycorrhizae had a strong effect on plant growth in undomesticated crops but had no effect on domesticated crops. When undomesticated plants were grown with mycorrhizae, they were 13% smaller aboveground (t=2.7, df=546, p<0.01, Fig 1A) and 21% smaller belowground (t=2.6, df=542, p<0.01, Fig 1B) than plants grown without mycorrhizae. On the other hand, domesticated plants grown with mycorrhizae were not substantially different aboveground (t=1.2, df=546, p=0.25, Fig 1A) or belowground (t=1.1, df=542, p=0.27, Fig 1B) compared to those grown without mycorrhizae. Overall, domesticated plants were 29% larger than undomesticated plants (F 1,503 =54, p<0.001), with domesticated eggplant, pepper, tobacco, and tomato growing larger than their undomesticated counterparts, while tomatillo and pepper grew to similar sizes to their wild counterparts (Supplemental Information 3). Domestication affected the flowering time of crops differently. Domesticated pepper, and tomato plants were more likely than their undomesticated versions to have flowered by the end of the experiment. Potato, chili and eggplant hadn’t flowered at all by the end of the experiment regardless of domestication status (Supplemental Information 3). Figure 1. The effect of domestication and mycorrhizae on A) belowground biomass and B) aboveground biomass. Mycorrhizae reduced biomass above and belowground, but only for domesticated plants (t=2.7, df=546, p<0.01). Bars represent model outputs +/- SE. Asterisks denote significance. Plant resistance traits Protease Inhibitors Similar to the effects on growth, mycorrhizae had an effect on undomesticated crops (z=1.96, p=0.0502), but had no effect on domesticated crops (z=0.45, p=0.65). Overall, domesticated plants had 25% lower protease inhibitor activity than undomesticated plants (F 1,234 =12, p<0.001, Fig. 2A). While there was a significant three-way interaction between mycorrhizae, crop, and domestication, none of the pairwise comparisons were significant (Supplemental Information 3). Protein Mycorrhizae had a marginal effect on the protein content of undomesticated crops (t.ratio=1.72, df=393, p=0.086), but had no effect on the protein content of domesticated crops (t.ratio=-0.82, df=393, p=0.41, Fig. 2B). The effects of domestication on leaf protein were dependent on crop (F 5,393 =4.9, p<0.001), with domestication increasing leaf protein in tobacco and decreasing it in eggplant (Supplemental Information 1). Figure 2. The effect of domestication and mycorrhizae on A) protease inhibitor activity as measured by percent inhibition and B) leaf protein. Bars represent model outputs +/- SE. Asterisks denote significance, while grey asterisks denote marginal significance. Trichomes The effect of domestication on mycorrhizae-conferred changes in trichome abundance was highly crop dependent, with mycorrhizae increasing total trichomes in domesticated tobacco, domesticated potato, and undomesticated potato, and decreasing trichomes in domesticated eggplant (Supplemental Information 3). Plant resistance to herbivores T. ni Consistent with the effects of protease inhibitors, mycorrhizae increased T. ni leaf area consumption in undomesticated plants (t=-2.3, df=106, p=0.022), but had no effect in domesticated plants (t=0.36, df=106, p=0.72 Fig. 3A) (interaction F 1,03 =8.4, p=0.005, Fig. 3A). But while the caterpillars fed more on mycorrhizal undomesticated plants, they did not gain more weight (t=-0.64, df=178, p=0.53, Fig 3B), indicating that these plants were a lower quality food source. In plants without mycorrhizae, T. ni that fed on domesticated plants ate twice as much as those that fed on undomesticated plants (F 1,106 =8.32, p=0.005). While there was a significant interaction between mycorrhizae and crop on weight gain (F 2,182 =3.2, p=0.043), there were no significant pairwise differences. Neither domestication (z-score=0.67, p=0.50) nor mycorrhizal status (z-score=0.64, p=0.52) of the host plant altered T. ni caterpillar survival. T. ni consumed more leaf area on plants that had higher levels of protein in the leaves (F 1,55 = 4.2, p=0.045). Neither protein nor protease inhibitors altered survival or weight gain (Supplemental Information 3). M. sexta The effects of domestication on mycorrhizae-conferred resistance to M. sexta leaf area consumed varied marginally by crop (F 5,395 =2.1, 0.06), although the effects of mycorrhizae were not significant in pairwise comparisons (Supplemental Information 3). Overall, M. sexta that fed on domesticated plants ate 20% more than those that fed on undomesticated plants (F 1,395 =7.3, 0.007). Mycorrhizae marginally increased M. sexta w eight gain in undomesticated plants (t=-1.9, df=346, p=0.064), but had no effect in domesticated plants (t=0.56, df=346, p=0.58). Neither protein nor protease inhibitor levels explained M. sexta leaf area consumed, or growth, but M. sexta had higher survival on plants with higher levels of protein (p=0.019) P. maculiventris Mycorrhizae suppressed P. maculiventris weight gain in undomesticated plants (t=2.1, df=139, p=0.036), but had no effect on P. maculiventris weight gain in domesticated plants (t=0.73, df=139, p=0.47, Fig. 3.3D). P. maculiventris gained the most weight on plants that had high levels of protein and low levels of trichomes (F 1,13 =6.3, p=0.027). There was a marginal interactive effect between protein and protease inhibitors such that P. maculiventris gained the most weight on plants that had high levels of protein and lower levels of protease inhibitors (F 1,14 =4.4, p=0.060). Figure 3. The effect of domestication and mycorrhizae on A) the amount of leaf tissue T. ni caterpillars consumed, B) T. ni percent weight gain C) M. sexta weight gain and D) T. ni percent weight change. Bars represent model outputs +/- SE. Asterisks denote significance, while grey asterisks denote marginal significance. Table 2 Mycorrhizae affected growth and resistance traits and resistance to herbivores in undomesticated plants, but had no effect in domesticated plants. The letter X and n.s. represent no effect, and not significant, respectively. Arrows indicate direction of effect. Italicized numbers are marginally significant. Undomesticated Plants Domesticated Plants Trait Effect of Mycorrhizae p value Effect of Mycorrhizae p value Growth Traits Aboveground biomass ↓ 0.008 X n.s. Belowground biomass ↓ 0.009 X n.s. Flowering time X n.s. X n.s. Resistance Traits Protease Inhibitors ↑ 0.0502 X n.s. Protein ↓ 0.089 X n.s. Trichomes X n.s. X n.s. Resistance to Herbivores T. ni leaf area consumed ↑ 0.022 X n.s. T. ni weight change X n.s. X n.s. T. ni survival X n.s. X n.s. M. sexta leaf area consumed X n.s. X n.s. M. sexta weight change ↑ 0.064 X n.s. M. sexta survival X n.s. X n.s. P. maculiventris weight change ↓ 0.036 X n.s. Discussion Domestication can alter plant-mutualist associations. However, key knowledge gaps remain in how domestication can alter mycorrhizae-conferred resistance to herbivores. Here we showed that a single mycorrhizal inoculum changed the growth and resistance of their undomesticated host plants (Fig 4) but had very little effect on their domesticated host plants. Domesticated plants had higher colonization than undomesticated plants. Surprisingly, we found that, in all but one species, domesticated plants had much higher levels of colonization than undomesticated plants. The process of domestication has greatly changed plant physiology, in general producing plants that are larger and less well defended against herbivores. Previous studies have shown that mycorrhizae preferentially colonize larger plants, likely due to the fact that larger plants have more carbon to give (Moora & Zobel, 1996; Weremijewicz et al. , 2016). Selection pressures for larger domesticated plants may lead to increased colonization. In general, both plants and mycorrhizae have the potential to regulate the symbiosis through differential sharing of resources (Kiers et al. , 2011). However, plant species differ in their ability to sanction non- beneficial mycorrhizae (Grman, 2012). Domesticated plants have lower expression of resistance traits. In our study, domesticated plants had lower levels of protease inhibitors. Pathways that control the expression of resistance traits including the jasmonate, salicylate and ethylene pathways, have also been implicated in regulating mycorrhizal colonization (Guinel & Geil, 2002; Herrera Medina et al. , 2003; Tejeda-Sartorius et al. , 2008; Herrera-Medina et al. , 2008; Martín-Rodríguez et al. , 2011). We speculate that changes in these pathways might reduce the plant’s ability to regulate colonization, resulting in the pattern we see of higher colonization despite the lack of benefits to the plant. These results contradict previous findings by Xing et al. (2012) and Turrini et al., (2016) which found that domesticated breadfruit and sunflowers had lower colonization than their undomesticated counterparts. Martin Robles et al (2018) also found lower colonization in domesticated crop plants, but the effect was quite variable across 27 crop species and the pattern was only evident at higher levels of fertilization. Broadly, the effects of domestication on growth and defense traits have been highly variable across plant families and crops (Pickersgill, 2007; Whitehead et al. , 2017). It seems that, like these other traits, the effects of domestication on mycorrhizal symbiosis may vary substantially across plant families and crops. Despite the higher colonization, we saw very little effect of mycorrhizae on growth or resistance in domesticated plants but significant effects in undomesticated plants. This disconnect between colonization levels and growth and resistance traits may mean that there is some reduction of symbiosis between plants and mycorrihizae. Our work highlights that measuring only colonization gives an incomplete snapshot of a complicated interaction. Future studies should consider measuring colonization, growth, and resistance traits to give a more complete picture of the interaction, and better help us understand how mycorrhizae fit into complex food webs in both natural and applied systems. Loss of resistance traits may have inadvertently reduced symbiosis with mycorrhizae Previous work by Martin Robles et al., (2018) found that domestication reduced the plant’s ability to associate with mycorrhizae, indicating some loss of symbiotic benefit for plants in these associations. Agriculture and domestication can disrupt the selection processes that govern the benefits of symbioses (Porter and Sachs, 2020), reducing selection and possibly resulting in an evolutionary loss of symbionts. The loss of symbiosis between mycorrhizae and domesticated plants has previously been attributed to fertilization regimes that might make associating with fungus unnecessary, as well as selective pressure to allocate carbon resources to growth instead of providing it to fungal symbionts. We suggest that the symbiosis may be further disrupted by changes in plant defensive chemistry and changes in pest pressure. Our results and previous studies found that domesticated plants have lower levels of defensive secondary metabolites and hormones (Whitehead et al. , 2017). We suggest that decreases in secondary metabolites and defensive chemistry may indirectly reduce the plant’s ability to associate with or properly sanction the fungus. It is well established that the jasmonate and salicylate pathways have a role in controlling both resistance traits and associations with fungal mutualists (Herrera Medina et al. , 2003; Tejeda-Sartorius et al. , 2008). By increasing pest control and actively selecting for plants with lower levels of unpalatable defenses, it is possible that humans have selected plants that are less well defended and also less able to associate with belowground mutualists like mycorrhizae. Mycorrhizae reduced plant biomass in undomesticated plants Our mycorrhizal inoculum reduced the above- and belowground biomass of their plant partners in undomesticated plants but not in domesticated plants. While there are ample studies that show a positive association between above- and belowground biomass of plants with mycorrhizae, our results provide evidence against the mutualistic paradigm that mycorrhizae always improve plant growth. Mycorrhizae have been known to reduce the biomass of their hosts (Stribley et al. , 1980; Wilson & Hartnett, 1997; Treseder, 2013; Stanescu & Maherali, 2017), ostensibly through the carbon cost (Olsson et al. , 2010). Plants can give up to 20% of their carbon-rich photosynthate to mycorrhizae (Hobbie, 2006), and this cost may outweigh the nutritional benefits they get in return. There is also ample evidence that mycorrhizae can suppress belowground biomass (Wurst et al. , 2004) and decrease the root to shoot ratio (Veresoglou et al. , 2012 ., and citations therein), although this is often attributed to the fact that mycorrhizae act as a secondary root system, reducing the need for the host plant to invest in belowground root tissue. In our system, both mechanisms may be at play as mycorrhizae suppressed growth belowground more dramatically than aboveground, but also significantly suppressed aboveground tissue. This result highlight that more investigation is needed before promoting mycorrhizae as tools for supporting plant growth and productivity in agricultural settings (Ryan & Graham, 2018). Mycorrhizae increased physical and chemical resistance traits and resistance to herbivores T. ni that fed on mycorrhizal undomesticated plants fed more but did not gain more weight. P. maculiventris that fed on these mycorrhizal undomesticated plants actually lost weight on average (Fig 3). In undomesticated plants, mycorrhizae suppressed M. sexta weight gain. We pose that, in undomesticated plants, mycorrhizae-conferred increases in protease inhibitors force T. ni to feed more to maintain their growth, and protease inhibitors and trichomes, or correlated traits, deter P. maculiventris from feeding. Mycorrhizae marginally decreased protein in undomesticated plants. Both T. ni and P. maculiventris also fed less on individual plants with lower levels of protein. This conforms to previous research showing that herbivores feed more when there are higher levels of nitrogen-rich protein in their diet, but it suggests that this is not a mechanism by which mycorrhizae alter herbivore performance. Our study suggests that mycorrhizae confer resistance in undomesticated plants through increasing chemical and physical defenses rather than through changing nutritional quality. However, we qualify these results with the reality that we used a single arbuscular inoculum, further research should investigate the variation in mycorrhizae to fully understand how plant resistance is modulated by symbiotic interactions in this system. Figure 4. In undomesticated plants, mycorrhizae increase protease inhibitors and trichomes but not protein. These adversely affected T. ni performance. Solid arrows represent significance. Red arrows represent a negative effect and black arrows indicate a positive effect. Our study builds on previous work by Xing et al., (2012) and Martin Robles et al., (2018) (see also Porter and Sachs, 2020). by demonstrating that, in addition to effects on growth and symbiosis, domestication has also led to changes in how mycorrhizae affect plant-herbivore interactions. In our study, mycorrhizae increased resistance to generalist T. ni and P. maculiventris and decreased resistance to specialist M. sexta . Consistent with our findings, a review by Koricheva et al., (2009) found that specialist (mono- and oligotrophic) chewing herbivores were more likely to benefit from mycorrhizae, while generalist chewing herbivores and piercing sucking herbivores were more likely to be negatively affected. Domestication reduced resistance traits and resistance to herbivores It is widely accepted that domesticated plants are larger and have less pest resistance than undomesticated plants (as reviewed by Whitehead et al., 2017). Our results support this, with domesticated plants growing larger, having lower defensive protease inhibitor activity, and increased feeding from T. ni and M. sexta . There is much debate as to the selective forces that have driven this lowered resistance. Potentially, selection against unpalatable defenses and selection for increased growth and yield both may result in less defended plants. Alternately, selection for more nutritious plants may have resulted in plants that are more attractive to herbivores. Our results lend credence to the hypothesis that selection for larger, less well defended plants has reduced resistance to herbivores, as our domesticated plants were larger and had lower levels of protease inhibitors but no difference in protein content. Consequences of mycorrhizal species As noted previously, our mycorrhizal species, Rhizophagus irregularis , is a commercially available species of mycorrhizal fungi used in agriculture and home gardens. Its highly generalist nature and well-documented use with solanaceous plants made it an ideal candidate for our study. However, we present these results with the caveat of species-specific traits for R. irregularis itself. This species is commercially produced for its ability to colonize well in many settings. However, traits that make it easily commercially available may also influence its interactions with our plants, as well as the ultimate outcomes we show in our study. Ultimately, future research should include a wider variety of mycorrhizal species, to fully understand how the suite of traits in different plants, mycorrhizae, and herbivores interact to create a more comprehensive insight into symbiotic and mutualistic interactions. Conclusion Our comparative approach of 6 domesticated and undomesticated solanaceous crop plants has shown that domestication has altered mycorrhizae-conferred resistance to herbivores and natural enemies. The lack of effect of our mycorrhizal inoculum in domesticated plants supports previous studies that find a breakdown in the plant-mycorrhizae mutualism in domesticated plants. These results, in combination with the finding that mycorrhizae suppressed plant growth, demonstrate the limitation of mycorrhizae as an agricultural tool. The disconnect between colonization and effects on growth and resistance demonstrates that measuring colonization only gives an incomplete snapshot of a complicated interaction. While we only used a single mycorrhizal inoculum, these results shed interesting insights on these interactions and highlight future work understanding the variation in mycorrhizae and whether there are consistent patterns in resistance and biomass traits in plants. Future studies should consider measuring colonization, growth, and resistance traits to give a more complete picture of the interaction, and better help us understand how mycorrhizae can benefit or harm plants. 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Keywords chemical ecology domestication multi-trophic mutualism mycorrhizae Authors Affiliations Zoe Getman-Pickering 0000-0002-5695-858X [email protected] University of Massachusetts Amherst View all articles by this author Jennifer Thaler Cornell University View all articles by this author Jules Davis 0000-0002-4902-675X University of Massachusetts Amherst View all articles by this author Heather Grab Penn State View all articles by this author Sheyla Finkner Sarepta Therapeutics Inc View all articles by this author Elyse McCormick University of Massachusetts Amherst View all articles by this author Abigail Getman-Pickering LeafByte View all articles by this author Kaitlin Deutsch University of Florida View all articles by this author Metrics & Citations Metrics Article Usage 141 views 63 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Zoe Getman-Pickering, Jennifer Thaler, Jules Davis, et al. 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