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Transgenerational effects of stress on reproduction strategy in the mixed mating plant Lamium amplexicaule

BMC Plant Biology volume 24, Article number: 794 (2024) Cite this article

Abstract

Background

The theory of Condition Dependent Sex predicts that – everything else being equal – less fit individuals would outcross at higher rates compared with fitter ones. Here we used the mixed mating plant Lamium amplexicaule, capable of producing both self-pollinating closed flowers (CL), alongside open flowers (CH) that allow cross pollination to test it. We investigated the effects of abiotic stress – salt solution irrigation – on the flowering patterns of plants and their offspring. We monitored several flowering and vegetative parameters, including the number and distribution of flowers, CH fraction, and plant size.

Results

We found that stressed plants show an increased tendency for self-pollination and a deficit in floral and vegetative development. However, when parentally primed, stressed plants show a milder response. Un-stressed offspring of stressed parents show reversed responses and exhibit an increased tendency to outcross, and improve floral and vegetative development.

Conclusions

In summary, we found that stress affects the reproduction strategy in the plants that experienced the stress and in subsequent offspring through F2 generation. Our results provide experimental evidence supporting a transgenerational extension to the theories of fitness associate sex and dispersal, where an individual’s tendency for sex and dispersal may depend on the stress experienced by its parents.

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Background

Mixed mating systems, allowing both self and cross reproduction, are common in plants [1, 2]. While both outcrossing and selfing result in the formation of new recombinant genotypes, outcrossing secures a greater variety of possible genetic combinations, and in the long run, increases adaptivity to varied environments in subsequent generations [3, 4]. For this reason, while growing under inferior conditions or stress, outcrossing with a different genotype could hold an advantage over selfing [5]. Moreover, outcrossing also involves pollen dispersal, potentially allowing the plant to disperse to a different environment [6]. On the other hand, outcrossing contributes less than self-fertilization to the transmission of parental genome [7], and depends on pollinators or nearby mates for successful reproduction [8,9,10,11]. Theories of fitness associated sex (FAS) and fitness associated dispersal (FAD) predict that individuals at mal condition should show a higher tendency for sex and dispersal compared with better adapted ones. Such condition-dependent variation can allow the genes responsible for plasticity in these behaviours to escape from maladapted genetic backgrounds, while staying linked with more adapted backgrounds [12,13,14,15,16], and may facilitate adaptation in the long term [17]. However, environmental factors such as stress are known to affect developmental and reproductive processes [18, 19] and might have a detrimental effect on pollination success of exposed individuals [20,21,22]. Stressful experiences can also influence plant future responses to stress, referred to as “stress memory” [23,24,25,26], and was reported to occur in plants at the seed stage [27, 28], and the adult plant stage, through epigenetic memory mechanisms [24, 29]. Heritability of priming, known as transgenerational stress memory [30,31,32], was shown in several studies on generation F1 [33, 34], and F2 [35], and shown to have phenotypic effects on the offspring’s leaf development, root : shoot ratio, and attractiveness to herbivores [36], generally enhancing offspring performance in stressful as well as benign conditions. This trend is especially noticeable in annual plants [37], inter alia, affecting reproductive floral traits such as flowering time [38].

In the following experiments, we used Lamium amplexicaule, a mixed mating annual species capable of high levels of self-reproduction through Cleistogamy [39]. A cleistogamous individual delivers heteroblastic inflorescences with dimorphic flower types (Fig. 1A): Chasmogamic flowers (CH) develop fully and produce big colourful petals, nectar, and ample pollen, while Cleistogamic flowers (CL) arrest petal development at an early stage [40], when the petals are concealing the gametes which continue to mature within the reduced closed structure, where self-pollination occurs [41].

Plants with dimorphic cleistogamy sense environmental factors before and during the flowering phase, and react by allocating buds’ development towards optimal morph [42, 43]. Producing higher rates of CH flowers, that are larger than CL flowers and rewarding, would increase the attractiveness of the plant to pollinators [44,45,46,47]. Moreover, the occurrence of many open flowers in a dense population can also generate a “magnet effect”, potentially benefitting not only the plants manifesting them, but also neighbouring plants, as pollinators are attracted to the site [48,49,50,51]. Inversely, producing higher rates of CL flowers would save resources since they are pronouncedly reduced in size [52]. The cleistogamic flower structure is associated with biotic [53] and abiotic [54, 55] stress tolerance, and in addition to genes controlling the flowering processes, some of the differently expressed genes in CL flowers tissue are involved in environmental stress responses [56, 57].

L. amplexicaule plants show plasticity in the rate of flower types among individuals, partially depending on the natural variation background [58], as well as external cues the plant detects: L. amplexicaule CH flowers percentages have been shown to increase with the intensity of light and supply of nutrients [59], in spring compared to autumn [60, 61], and with long photoperiod and high temperatures [62]. The effect of competition was mixed: CH fraction increased in unfavourable environments with high density [63], but decreased under interspecific competition [64]. In addition, unfavourable environments with high density reduced overall flowering (both CH and CL) [63], and interspecific competition decreased above ground biomass [65].

The transgenerational effects of environmental conditions on the reproduction strategy of L. amplexicaule plants haven’t been tested yet to our knowledge, and the goal of this study is to extend our understanding of these effects on the following generations, focusing on the question: will salt stress conditions lead to an increase in the likelihood of outcrossing, within or between generations?

Since plants evolved to withstand continuous environmental stress, we hypothesize the diminishing effects of salt stress will be reduced on the second generation of stress exposure, and reversed in offspring growing in benign conditions, revealing increased outcrossing rates, either directly with CH flowers, or indirectly by better advertisement to pollinators.

Materials and methods

Sampling seeds and creating single seed descent lines

Lamium amplexicaule seeds were identified and sampled in spring (March, temp range 21° / 10°, 5 rain days on average per month) from individuals of a natural population in Netzer Sereni, Israel (31°55′33.96″N 34°49′46.2″E), by Nir Ohad, Lilach Hadany, and Ariel Gueijman. Seeds were taken directly from the sepals after the senesces of the flowers, so the type of flower (CH\CL) is unknown. L. amplexicaule seeds are characterized by endogenous dormancy [66], so seeds were given a hibernation period of nine months, before placement in damp planting soil in standard germination trays (4 cm diameter x 4.5 cm height round pots). Sprouts were transferred to individual pots (9 cm x 9 cm x 10 cm) containing planting soil, and grown separately under control conditions: 16 L\8D, 21–23℃, irrigated with water (H2O), fluorescent light combining white, blue, and purple bulbs (Fluora, Daylight, and Cool White lamps by OSRAM), in a controlled growth room with no pollinator activity, meaning subsequent seeds were the result of self-pollinating in both flower types (CH/CL). Individual seeds were collected at senescence. After a hibernation period of minimum nine months, sibling seeds were germinated and grown in control conditions, separately for each founder lineage, to prevent any minor possibility of cross pollination between lines. Subsequent seeds of sibling plants were collected combined as Single Seed Descent (SSD) line pool. The experiments presented here were performed on two SSD lines to investigate reproduction strategies: line 15 and 8, as indicated below. A voucher specimen of the material used in this study has been deposited in the Israel Plant Gene Bank (https://igb.agri.gov.il/web/) a publicly available herbarium with the following deposition number: 10,029,218.

Experiment 1

In July 2013 seeds of SSD line 15 were germinated and divided into two irrigation regimens: control plants- (denoted by C) irrigated with water (H2O), and stressed plants (S)- irrigated with 50 mM NaCl solution (Fig. 1B).

We chose to use salt irrigation treatment (50 mM NaCl) to inflict stress for a few reasons: first, it is easy to quantify and apply, and allows other growth conditions to stay stable, and equal, for all groups. Second, salinity of soil is a feasible type of stress L. amplexicaule plants may face in Mediterranean habitats, having deleterious effects on yield and plant physiology [67]. Concentration of 50 mM NaCl was used based on preliminary tests suggesting this salinity level allows plants to survive and reproduce.

Flowering patterns were not measured in these plants. Both groups were constantly held under fluorescent light combining white, blue, and purple bulbs (Fluora, Daylight, and Cool White lamps by OSRAM), 16 L\8D, 21–23℃. The irrigation solution (H2O / NaCl) was constantly applied during the plants’ life cycle (up to four months). We determined this generation as P, since past cycles were under benign conditions prior to this experiment. Seeds were collected and stored as population pool for group C and group S. On August 2015 100 seeds of control lineage and 100 seeds of stress lineage were sewn in customized germination trays. After two weeks, 64 seedlings of each lineage were randomly chosen and transferred to individual pots (9 cm x 9 cm x 10 cm), and divided again according to past and present generation growth conditions (Fig. 1B): CC, CS, SC, and SS, accounting a total of 32 plants for each group. Individual pots were individually wrapped in a clear organza net bag for two purposes: preventing tangling of stalks, and collecting individual seeds. Pots were placed in trays, each tray containing 8 plants of the same group, and placed on shelves in a growth room. Two days after pot transfer the irrigation regime began: CS and SS plants were irrigated with 50 mM NaCl solution, while CC and SC were irrigated with H2O, throughout the entire plant lifespan. The first flowers began to appear on September 2015. The main stalk of each plant was measured as it originates from the hypocotyl and therefore easily traceable. Flowers on the main stalks were counted every 2 to 3 days, indicating date of count, whorl position, and amount of CL and CH flowers. The count was stopped on December 2015 as the last plants began to senesce. The main stalk of each plant was measured in cm. During the experiment two plants were eliminated due to dehydration and final groups sizes analysed were: CS n = 32, SS n = 32, SC n = 31, and CC n = 31.

Experiment 2

In February 2014 single seed descendants of line 8 were formed as described in SSD 15, to form lineages CC, CS, SC, and SS. On May 2016, seeds of each lineage were sewn in customized germination trays. After 3 weeks, 32 seedlings from each lineage were randomly chosen and transferred to individual pots (9 cm x 9 cm x 10 cm), to create study groups CCC, CSC, SSC, and SCC. Plants were watered with H2O throughout their entire life cycle. Plants were handled and counted in the same manner as in experiment 1. The flower count began on June 2016 and ended on 19.12.2016. During the experiment eleven plants were eliminated due to dehydration and final groups sizes analysed were: CSC n = 32, SSC n = 30, SCC n = 32, and CCC n = 23.

Parameters of comparison and statistical analysis

to assess plant reproduction we collected data for comparisons from the main stalk of each study plant. We monitored the total number of flowers, the percentage of CH flowers, number of flowering whorls, average number of flowers in each whorl, and plant height. In light of non-normal distributions of these parameters within treatment populations, we chose conservative nonparametric tests to conduct comparisons: Mann-Whitney Wilcoxon U test for averages [68], F test for variance [69]. P-values of each analysis were corrected using the Benjamini-Hochberg FDR procedure for multiple comparisons [70]. Experimental data for each SSD line were analysed separately. Since CH rates are sensitive to external cues, we compared groups of SSDs grown simultaneously in the same growth room.

Results

In the experiments presented here, we tested the effect of soil salinity stress over one and two consecutive generations, on the stressed individuals and their F1 and F2 offspring as compared with control individuals, using two Single Seed Descendant (SSD) lines (Fig. 1B). We monitored several flowering and vegetative characteristics, including the number and distribution of flowers, CH fraction, and plant size, and tested whether these characteristics were altered in response to stress, either in the stressed generation or in their offspring.

Fig. 1
figure 1

(A): L. amplexicaule dimorphic flowers. CL performs self-pollination. CH performs both self and cross pollination. (B): Schematic illustration depicting the experimental regime. Single seed descendants were divided into two groups- one watered with 50 mM NaCl solution (S, red arrow), and the other watered with H2O (C, blue arrow). Seeds collected were divided in the same manner, forming a total of four study groups. Subsequent seeds were all irrigated with H2O. Group names are composed of chronological C or S indicating irrigation type of each generation. Experiment 1 included groups of SSD line 15: plants experiencing ongoing salt stress for one generation (CS), two generations (SS), non-stress F1 offspring (SC), and control (CC). Experiment 2 included groups of SSD line 8: non-stress F1 offspring of one stress generation (CSC), and two stress generations (SSC), non-stress F2 offspring of one stress generation (SCC), and control (CCC). We define the offspring of stressed parents (groups SS, SC, CSC, SSC, and SCC) as stress-primed, and the offspring of unstressed parents (CC, CS, CCC) as unprimed

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Reproduction strategy - rate of Chasmogamy (CH)

Plants of SSD line 15 that encountered salt stress for the first time (CS) were almost completely cleistogamous. The average rate of CH flowers was only 0.5% in this group, while the control rate was 19% (P < 10E-10). However, in plants that experienced stress for two subsequent generations (SS) the average increased to 17% (P < 0.0011 compared to CS), still significantly lower than control (P < 0.0072) (Fig. 2A). Interestingly, SS plants (under two generations of stress) showed a bimodal distribution of CH rates (Fig. 2B, top). While many individuals produced ~ 0% CH flowers (23 individuals out of 32), a considerable minority (8 individuals out of 32) produced 50 − 80% CH flowers. Such high rates have never been observed in any of our control populations. Non-stressed F1 plants (SC) of this line produced 18% CH flowers, not different from the control rate of 19%.

In the second experiment done on SSD line 8, the control average for CH rates was 14%. Unlike line 15, the unstressed offspring of stressed parents of line 8 exceeded the corresponding control average. Among F1 populations, CSC plants measured 34% (P < 10E-4), and SSC reached 43%, significantly higher compared to control (P < 10E-5), as well as to CSC plants (P < 0.036), suggesting an accumulating response (Fig. 2B, bottom). In F2 SCC plants, CH rate averaged 35%, significantly higher than control as well (P < 10E-6) (Fig. 2A).

Fig. 2
figure 2

(A): Box-plots for CH (chasmogamy) rates. Group means (x). Significance according to Mann- Whitney U test corrected under Benjamini-Hochberg FDR procedure for multiple comparisons, done separately for each SSD line. Each bar is the mean value ± SD. Different letters indicate significant differences among groups. (B): Frequency of CH rate within populations of line 15 (top) and line 8 (bottom). X-axis: CH fraction. Y-axis: frequency in the population

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The effect of stress on other vegetative and reproductive traits

In addition to alterations in CH rates, other floral characteristics affecting pollination were significantly altered in the treatment groups. In line 15, CC plants had an average of 109 total flowers, average whorl size of 8 flowers, and average hight of 40 cm (Fig. 3). Vegetative and reproductive characteristics were reduced under stress, as both CS and SS plants bloomed less, averaging 41 and 42 total flowers (P < 10E-10 compared to CC in both cases) (Fig. 4A). In addition, they were shorter in height compared to control, reaching 29 cm (P < 10E-10), and 32 cm (P < 10E-08) (Figs. 3 and 4B). However, parental priming moderated some of the effects of stress: while in CS the number of flowers per whorl decreased to 6.5 (P < 10E-08 compared to CC), SS average increased to 8 flowers per whorl (P < 0.002 compared to CS), not different from the control average (Fig. 4C). In addition, SS plants were taller than CS plants (P < 0.0012) (Fig. 4B). F1 offspring (SC) of this line had increased overall flowering compared to control, with an average of 128 flowers (P < 0.022), whorl containment of 9.5 flowers per whorl (P < 0.012), and increased height, averaging 50 cm (P < 0.0002) (Figs. 3 and 4).

Fig. 3
figure 3

Schematic illustration depicting the average main stalk of line 15 study groups, proportioned to plant height and number of whorls. The bar plot represents the distribution of flowers upon whorls. Grey: Cleistogamic (CL, closed) flowers. Purple: Chasmogamic (CH, open) flowers. Under stalks is an illustration of the average whorl containment (flowers per whorl). Parentally primed plants (SS, SC) produced more flowers with higher CH rates in the lower whorl positions forming earlier. Group sizes: Line 15: CS n = 32, SS n = 32, SC n = 31, and CC n = 31. Line 8: CSC n = 32, SSC n = 30, SCC n = 32, and CCC n = 23

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In the second SSD examined, line 8, we found that control plants (CCC) averaged 98 total flowers, 7 flowers per whorl, and 28 cm in height. In general, unstressed F1 and F2 offspring of stressed plants grew and flowered more vigorously than control plants: F1 CSC plants had 110 flowers (ns), 9 flowers per whorl (P < 0.0031), and were 27 cm tall (ns). F1 SSC plants had 133 flowers (P < 0.0132), 10 flowers per whorl, significantly higher than control (P < 10E-5) and CSC plants (P < 0.0132), and were 29 cm tall (ns). F2 SCC plants had 125 flowers (P < 0.0329), 10 flowers per whorl (P < 10E-5), and were 33 cm tall (ns) (Fig. 4).

Fig. 4
figure 4

Box-plots for (A): number of flowers, (B): plant height, (C): number of flowers per whorl. Group means (x). Significance according to Mann- Whitney U test corrected under Benjamini- Hochberg FDR procedure for multiple comparison, done separately for each SSD line. Each bar is the mean value ± SD. Different letters indicate significant differences among groups

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Spatial and temporal distribution of flowers

Under control conditions, plants displayed mostly cleistogamic (CL) flowers in lower whorls, while chasmogamous (CH) flowers developed later in the season, on higher whorls (Fig. 3). We tested if differences in reproductive strategy are significant at the first two flowering whorl positions, formed at the very beginning of the reproductive phase. In line 15, the control averaged 11.7 flowers, with 1.9% CH. Under exposure to stress, CS plants produced 10.6 flowers (P < 0.02) with 0.31% CH (ns).

Parental exposure to stress resulted in increased investment in the lower whorls of the offspring, regardless to present generation conditions: SS plants had 14.5 flowers, significantly higher compared to control plants (P < 0.0031), and CS plants (P < 10E-3), with a higher average of CH of 16% (P < 0.0768 compared with CC, P < 0.0466 compared with CS). SC plants averaged 14.5 flowers (P < 0.0072 compared to CC) with 6.9% CH flowers (ns, P = 0.2041).

In line 8, the control average in the first two whorls was 11.4 flowers, with 2.6% CH rate. Unstressed F1 and F2 offspring produced more flowers in the first two whorls: CSC produced 13.3 (ns), SSC produced 15.4 (P < 0.0004), and SCC produced 18.9 (P < 10E-5), with higher CH rates as well: CSC averaged 8% (P < 0.02), SSC 18.9% (P < 10E-3), and SCC 18% (P < 0.0002).

Discussion

In this study, we have tested L. amplexicaule reproductive strategies over generations following stress conditions. We found that stress has conflicting effects on the stressed individuals and their offspring (Table 1). Under stress, CH rate decreased, and stressed plants produced much fewer flowers in total, had fewer flowers per whorl, and were shorter in height (Figs. 3 and 4). However, stressed offspring of stressed plants partially overcame the stress, with increased average and variance of CH rate (Figs. 2 and 3), plant height (Fig. 4B), and number of flowers per whorl (Figs. 3 and 4C). Unstressed F1 offspring of stressed parents showed an increased number of total flowers and flowers per whorl (in both lines, Fig. 4A, C), increased CH rate in F1 and F2 (in line 8, Fig. 2), and increased height of F1 plants (in line 15, Figs. 3 and 4B).

Table 1 Treatment groups trend of response as compared to corresponding control for the following parameters: rate of Chasmogamic flowers (%CH), total number of flowers, number of flowers per whorl (F/W), and plant height
Full size table

Our experiments were performed in a controlled environment, discounting factors occurring in wild populations: First, our experiments included only plants that germinated at the first irrigation event (see methods), excluding hibernator seeds [71] that might have shown a different response. Second, due to the lack of pollinators in our growth room, no actual cross pollination occurred in CH flowers, and seeds produced by both flower types were the result of self-pollination. Successful pollination events might be another factor affecting the chasmogamy rates of an individual and their offspring [72, 73]. Third, plants were grown in individual pots and did not share soil. Chasmogamy rate is possibly affected by the presence of neighbors as potential mates or competitors, in addition to considerations of density-dependent gene dispersal [74, 75]. Fourth, CH rates may fluctuate substantially among individuals and populations of cleistogamic plant species [76,77,78,79,80], and specifically among L. amplexicaule populations [60, 81], which may explain inconsistency between SSDs. The above should be considered when extrapolating these results to the expected response of wild populations in natural environment. Nevertheless, we provide evidence of transgenerational stress response of mixed mating individuals.

On one hand, as predicted by condition dependent sex theory [12,13,14,15,16], the advantages of outcrossing are higher for stressed plants, thus CH rates are expected to increase. On the other hand, stress is known to cause deficiencies in vegetative growth and yield [82,83,84], and may decrease the ability of production, and later on the effectiveness, of CH flowers. Our results reflect both factors. Un-primed stressed plants (CS) showed restricted growth and decreased flowering, particularly costly CH flowers, while the increased CH rates and earlier temporal production of CH flowers in parentally primed stressed plants (SS, Fig. 2) suggests that chasmogamy is not determined only by energy limitations [85, 86], and parental experience can contribute to the reproductive strategy of the offspring, and may allow condition-dependent sex to be expressed in the generation following the stress exposure.

Evidence of transgenerational plasticity has been reported in angiosperms [87,88,89], suggesting parental effects may override immediate adaptive responses to stress and increase variation in offspring [90]. Indeed, we found the variance of SS plants had increased to form two distinct sub populations (Fig. 2B, top), with some individuals strictly self-pollinating and ensuring reproduction, while others cross-pollinate at very high rates, increasing the chances of their offspring to be heterozygous or move to a different environment through pollen transmission.

Our results provide further information about transgenerational effects in a mixed mating plant L. amplexicaul which allows for monitoring of reproduction strategy under stress. Parental priming was noticeable when increased CH rates were found in stress offspring grown under benign conditions (line 8, Fig. 2), suggesting the levels of response may be accumulating according to the duration of stress exposure (SSC were more affected than CSC). Such accumulating responses have been reported in Arabidopsis thaliana plants, as multiple-generation exposure to salt stress differentially impacted F2 offspring traits compared to single-generation exposure [38], and under prolonged heat stress, plants exhibited higher tolerance and frequency of homologous recombination, correlated with the duration of exposure [91].

In addition, whether grown in benign conditions or under stress, parentally primed plants whorl containment (flowers per whorl) had significantly increased (Fig. 4C), which could contribute to pollinator attraction [44, 49, 92, 93] and facilitate outcrossing.

Another interesting result regarding the reproductive success of non-stressed offspring of stress parents is that F2 plants that experienced stress once (SCC) performed better than F1 plants that experienced stress once (CSC) (Fig. 4B, C). Abiotic stress has been reported to adversely affect the quality and yield of seeds and fruit growth [94], reduce seeds vigour, mass, and germination [94, 95], and lead to less intense primary seed dormancy [96]. Such traits may correlate with the juvenile plant performance [97, 98], and may impact the adult stage [99]. The reduced effect observed in CSC (F1) compared to SCC (F2) can possibly be explained by the fact that F1 generation was subjected to stress during its embryonic development, which may override the beneficial effects of FAS to increase outcrossing. Thus, any response observed in generation F1 reproduction strategy might have been determined at the seed stage. Conversely, F2 did not encounter stress at any stage, thus the observed effects of parental stress are most likely inherited, rather than environmentally induced. Since L. amplexicaule plants in these experiments are of SSD lines, phenotypic differences among treatment groups within lines may have arisen at least in part from epigenetic regulation. Epigenetic changes due to stress exposure were found to be inherited in multiple studies [100,101,102,103], and in some cases to increase the resistance of progeny to the same stressor [104,105,106], as well as cross stressors [31, 100]. In addition to plants, transgenerational inheritance of stress memory has been found in animal models—Drosophila [107] and C. elegans [108, 109]. Our results provide further evidence for transgenerational inheritance of stress memory, and introduce a connection between parental stress and offspring mating strategy in plants.

Conclusions

In summary, we found that stress has an effect on L. amplexicaule reproduction strategy in plants exposed to stress and in subsequent offspring through F2 generation. Increased frequency of sex and dispersal in response to stress have been demonstrated both empirically [110,111,112,113] and theoretically [13]. Our work is consistent with this line of research but adds a significant component: that plasticity in variation (e.g. increased sex and dispersal) can occur not in the stressed individuals themselves, but rather in their offspring. Our results thus support an extension of condition dependent sex theory to transgenerational effects. In incidences where stressed plants lack the resources to achieve effective cross-pollination, F1 and F2 may increase outcrossing and dispersal efforts as a response. We suggest that a second-generation increase in variation in response to stress may be common in organisms where the stress increases the costs of sex and dispersal, as is the case for many plants.

Data availability

The datasets used and/or analyzed during the current study are available from the corresponding author upon request.

Abbreviations

CL:

cleistogamic flower- performs self-pollination

CH:

Chasmogamic flower- performs self and cross pollination

SSD:

single seed decent

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Acknowledgements

We would like to dedicate this manuscript in honor of the departed Prof. Dan Eisikowitch, a pioneer in the field of plant pollination, who introduced us to our first steps with Lamium plants. We thank Ariel Gueijman who originally collected seeds from a wild population to create the single seed descendant lines studied in these experiments. We thank Jenia Smoliarenko and Shirley Croitoru for their assistance in handling plants and assisting in phenotyping.

Funding

these researches were supported in part by the Israel Science Foundation Grants No. 1714/14 and 2064/18.

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  1. Lilach Hadany and Nir Ohad contributed equally to the research.

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  1. Department of Molecular Biology and Ecology of Plants, Tel Aviv University, Tel Aviv, 6997801, Israel

    Mor Binder, Eyal Zinger, Lilach Hadany & Nir Ohad

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  1. Mor Binder
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  2. Eyal Zinger
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LH and NO conceived the research. MB, EZ, LH, and NO designed the study. MB compiled and analyzed data. MB, LH and NO wrote the manuscript with all coauthors contributing to revisions.

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Correspondence to Nir Ohad.

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Binder, M., Zinger, E., Hadany, L. et al. Transgenerational effects of stress on reproduction strategy in the mixed mating plant Lamium amplexicaule. BMC Plant Biol 24, 794 (2024). https://doi.org/10.1186/s12870-024-05458-x

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