Flavonoids are specialized metabolites largely widespread in plants where they play numerous roles including defense and signaling under stress conditions. These compounds encompass several chemical subgroups such as flavonols which are one the most represented classes. The most studied flavonols are kaempferol, quercetin and myricetin to which research attributes antioxidative properties and a potential role in UV-defense through UV-screening mechanisms making them critical for plant adaptation to climate change. Despite the great interest in flavonol functions in the last decades, some functional aspects remain under debate. This review summarizes the importance of flavonoids in plant defense against climate stressors and as signal molecules with a focus on flavonols in Mediterranean plant species. The review emphasizes the relationship between flavonol location (at the organ, tissue and cellular scales) and their function as defense metabolites against climate-related stresses. It also provides evidence that biosynthesis of flavonols, or flavonoids as a whole, could be a crucial process allowing plants to adapt to climate change, especially in the Mediterranean area which is considered as one of the most sensitive regions to climate change over the globe.

Conifer forests worldwide are becoming increasingly vulnerable to the effects of climate change. Although the production of phenolic compounds (PCs) has been shown to be modulated by biotic and abiotic stresses, the genetic basis underlying the variation in their constitutive production level remains poorly documented in conifers. We used QTL mapping and RNA-Seq to explore the complex polygenic network underlying the constitutive production of PCs in a white spruce (Picea glauca) full-sib family for 2 years. QTL detection was performed for nine PCs and differentially expressed genes (DEGs) were identified between individuals with high and low PC contents for five PCs exhibiting stable QTLs across time. A total of 17 QTLs were detected for eight metabolites, including one major QTL explaining up to 91.3% of the neolignan-2 variance. The RNA-Seq analysis highlighted 50 DEGs associated with phenylpropanoid biosynthesis, several key transcription factors, and a subset of 137 genes showing opposite expression patterns in individuals with high levels of the flavonoids gallocatechin and taxifolin glucoside. A total of 19 DEGs co-localized with QTLs. Our findings represent a significant step toward resolving the genomic architecture of PC production in spruce and facilitate the functional characterization of genes and transcriptional networks responsible for differences in constitutive production of PCs in conifers.

Increasing aridity in the Mediterranean region will result in longer and recurrent drought. These changes could strongly modify plant defenses, endangering tree survival. We investigate the response of chemical defenses from central and specialized metabolism in Quercus pubescens Willd. to future Mediterranean drought using a long-term drought experiment in natura where trees have been submitted to amplified drought (~ –30% annual precipitation) since April 2012. We focused on leaf metabolites including chlorophylls and carotenoids (central metabolism) and flavonols (specialized metabolism). Measurements were performed in summer from 2016 to 2022. Amplified drought led to higher concentrations of total photosynthetic pigments over the 2016–2022 period. However, it also led to lower AZ/VAZ and flavonol concentrations. Additionally, chemical defenses of Q. pubescens responded to previous precipitation where low precipitation 1 year and/or 2 years preceding sampling was associated to low concentrations of VAZ, flavonol and high neoxanthin concentrations. Our study indicates that the decline of flavonol concentration under long-term drought is counterbalanced by a higher production of several central metabolites. Such results are potentially due to an adjustment in tree metabolism, highlighting the importance of performing long-term experimental studies in natura for assessing drought legacy effects and thus forest adaptation to climate change.

In the Mediterranean region, a reduction of annual precipitation and a longer and drier summer season are expected with climate change by the end of the century, eventually endangering forest survival. To cope with such rapid changes, trees may modulate their morpho-anatomical and physiological traits. In the present study, we focused on the variation in leaf gas exchange and different leaf morpho-anatomical functional traits of Quercus pubescens Willd. in summer using a long-term drought experiment in natura consisting of a dynamic rainfall exclusion system where trees have been submitted to amplified drought (AD) (~−30% of annual precipitation) since April 2012 and compared them with trees under natural drought (ND) in a Mediterranean forest. During the study, we analyzed net CO₂ assimilation (A_n), stomatal conductance (g_s), transpiration (E), water-use efficiency (WUE), stomatal size and density, density of glandular trichomes and non-glandular trichomes, thickness of the different leaf tissues, specific leaf area and leaf surface. Under AD, tree functioning was slightly impacted, since only A_n exhibited a 49% drop, while g_s, E and WUE remained stable. The decrease in A_n under AD was regulated by concomitant lower stomatal density and reduced leaf thickness. Trees under AD also featured leaves with a higher non-glandular trichome density and a lower glandular trichome density compared with ND, which simultaneously limits transpiration and production costs. This study points out that Q. pubescens exhibits adjustments of leaf morpho-anatomical traits which can help trees to acclimate to AD scenarios as those expected in the future in the Mediterranean region.

The intensity, frequency and severity of drought events are projected to increase in the Mediterranean region under ongoing climate change. This may impact plant growth and survival in Mediterranean forests leading to altered ecosystem dynamics. The functional diversity of soil macrofauna may have a critical role in mitigating negative climate change effects as it determines soil structure and regulates biogeochemical cycling as well as water retention. However, very little is known on how changes in climate and soil biodiversity conjointly influence plant growth. In a 5‐year experiment, we tested two levels of soil macrofauna functional diversity (Low and High), using combinations of endogeic and anecic earthworms, diplopods and isopods, and measured growth and biomass allocation in three tree species ( Acer monspessulanum , Arbutus unedo and Quercus pubescens ) as well as total tree biomass production. Our results showed that tree growth was overall only moderately affected by stronger drought and functional diversity of soil macrofauna. The effects of functional diversity of soil macrofauna differed among tree species. High functional diversity was associated with greater stem diameter in Q. pubescens . In contrast, low functional diversity was associated with greater stem diameter in A. unedo and proportionally greater biomass allocation to roots in A. monspessulanum . Individual macrofauna groups had stronger effects than overall functional diversity. Endogeic earthworm presence was associated with greater ecosystem‐level fine root and total plant biomass, but with lower fine root and total biomass in A. unedo . Diplopod presence was associated with greater stem diameter and higher total tree biomass in Q. pubescens . Overall, four consecutive summers of intensified drought had few significant effects on tree growth and biomass production, possibly because of high variability among individual trees. Most macrofauna effects were independent of drought, although drought modified the effect of diplopods on ecosystem‐level shoot biomass. These results show that macrofauna effects on tree growth depend on both macrofauna group and tree species identity. Read the free Plain Language Summary for this article on the Journal blog.