An Agronomic Approach to Pine Nut Production by Grafting Stone Pine on Two Rootstocks

Stone pine (Pinus pinea) grows in natural stands within the Mediterranean basin and its nut is highly appreciated for its nutritional profile. Nevertheless, a decline in this species due to biotic and abiotic damages is currently being clearly observed. This situation has led to its development as a nut crop, to try to save its production and obtain regular harvests over the years. Under this agronomic scenario, the aim of this work was to compare the behavior of the stone pine grafted onto two rootstocks, P. pinea (PP) and P. halepensis (PH), by evaluating cone productivity, tree growth response, mast seeding patterns and pine nut composition. The field test was composed of 14 PH and 14 PP, randomly distributed into groups of 4–5 trees/rootstock. Data were from seven productive growing seasons. The results show higher growth and ripe cone production on PP rather than PH, although the productivity (cones/m2 canopy) was similar. Any effect of rootstock was observed on the mast seeding pattern and weight of cones, while the pine nut composition showed differences in the fatty acids content. The global quality of production was similar in PH and PP.

Studying the genetic basis of masting

The mechanisms underlying mast seeding have traditionally been studied by collecting long-term observational data on seed crops and correlating seedfall with environmental variables. Significant progress in ecological genomics will improve our understanding of the evolution of masting by clarifying the genetic basis of masting traits and the role of natural selection in shaping those traits. Here, we summarize three important aspects in studying the evolution of masting at the genetic level: which traits govern masting, whether those traits are genetically regulated, and which taxa show wide variation in these traits. We then introduce recent studies on the molecular mechanisms of masting. Those studies measure seasonal changes in gene expression in natural conditions to quantify how multiple environmental factors combine to regulate floral initiation, which in many masting plant species is the single largest contributor to among-year variation in seed crops. We show that Fagaceae offers exceptional opportunities for evolutionary investigations because of its diversity at both the phenotypic and genetic levels and existing documented genome sequences. This article is part of the theme issue ‘The ecology and evolution of synchronized seed production in plants’.

The effects of ENSO and the North American monsoon on mast seeding in two Rocky Mountain conifer species

We aimed to disentangle the patterns of synchronous and variable cone production (i.e. masting) and its relationship to climate in two conifer species native to dry forests of western North America. We used cone abscission scars to reconstruct ca 15 years of recent cone production in Pinus edulis and Pinus ponderosa , and used redundancy analysis to relate time series of annual cone production to climate indices describing the North American monsoon and the El Niño Southern Oscillation (ENSO). We show that the sensitivity to climate and resulting synchrony in cone production varies substantially between species. Cone production among populations of P. edulis was much more spatially synchronous and more closely related to large-scale modes of climate variability than among populations of P. ponderosa . Large-scale synchrony in P. edulis cone production was associated with the North American monsoon and we identified a dipole pattern of regional cone production associated with ENSO phase. In P. ponderosa , these climate indices were not strongly associated with cone production, resulting in asynchronous masting patterns among populations. This study helps frame our understanding of mast seeding as a life-history strategy and has implications for our ability to forecast mast years in these species. This article is part of the theme issue ‘The ecology and evolution of synchronized seed production in plants’.

Climate change and plant reproduction: trends and drivers of mast seeding change

Climate change is reshaping global vegetation through its impacts on plant mortality, but recruitment creates the next generation of plants and will determine the structure and composition of future communities. Recruitment depends on mean seed production, but also on the interannual variability and among-plant synchrony in seed production, the phenomenon known as mast seeding. Thus, predicting the long-term response of global vegetation dynamics to climate change requires understanding the response of masting to changing climate. Recently, data and methods have become available allowing the first assessments of long-term changes in masting. Reviewing the literature, we evaluate evidence for a fingerprint of climate change on mast seeding and discuss the drivers and impacts of these changes. We divide our discussion into the main characteristics of mast seeding: interannual variation, synchrony, temporal autocorrelation and mast frequency. Data indicate that masting patterns are changing but the direction of that change varies, likely reflecting the diversity of proximate factors underlying masting across taxa. Experiments to understand the proximate mechanisms underlying masting, in combination with the analysis of long-term datasets, will enable us to understand this observed variability in the response of masting. This will allow us to predict future shifts in masting patterns, and consequently ecosystem impacts of climate change via its impacts on masting. This article is part of the theme issue ‘The ecology and evolution of synchronized seed production in plants’.

Mast seeding promotes evolution of scatter-hoarding

Many plant species worldwide are dispersed by scatter-hoarding granivores: animals that hide seeds in numerous, small caches for future consumption. Yet, the evolution of scatter-hoarding is difficult to explain because undefended caches are at high risk of pilferage. Previous models have attempted to solve this problem by giving cache owners large advantages in cache recovery, by kin selection, or by introducing reciprocal pilferage of ‘shared’ seed resources. However, the role of environmental variability has been so far overlooked in this context. One important form of such variability is masting, which is displayed by many plant species dispersed by scatterhoarders. We use a mathematical model to investigate the influence of masting on the evolution of scatter-hoarding. The model accounts for periodically varying annual seed fall, caching and pilfering behaviour, and the demography of scatterhoarders. The parameter values are based mostly on research on European beech ( Fagus sylvatica ) and yellow-necked mice ( Apodemus flavicollis ). Starvation of scatterhoarders between mast years decreases the population density that enters masting events, which leads to reduced seed pilferage. Satiation of scatterhoarders during mast events lowers the reproductive cost of caching (i.e. the cost of caching for the future rather than using seeds for current reproduction). These reductions promote the evolution of scatter-hoarding behaviour especially when interannual variation in seed fall and the period between masting events are large. This article is part of the theme issue ‘The ecology and evolution of synchronized seed production in plants’.

An assessment of temporal variability in mast seeding of North American Pinaceae

Our overall objective is to synthesize mast-seeding data on North American Pinaceae to detect characteristic features of reproduction (i.e. development cycle length, serotiny, dispersal agents), and test for patterns in temporal variation based on weather variables. We use a large dataset ( n = 286 time series; mean length = 18.9 years) on crop sizes in four conifer genera ( Abies , Picea , Pinus , Tsuga ) collected between 1960 and 2014. Temporal variability in mast seeding (CVp) for 2 year genera ( Abies , Picea , Tsuga ) was higher than for Pinus (3 year), and serotinous species had lower CVp than non-serotinous species; there were no relationships of CVp with elevation or latitude. There was no difference in family-wide CVp across four tree regions of North America. Across all genera, July temperature differences between bud initiation and the prior year (Δ T ) was more strongly associated with reproduction than absolute temperature. Both CVp and Δ T remained steady over time, while absolute temperature increased by 0.09°C per decade. Our use of the Δ T model included a modification for Pinus , which initiates cone primordia 2 years before seedfall, as opposed to 1 year. These findings have implications for how mast-seeding patterns may change with future increases in temperature, and the adaptive benefits of mast seeding. This article is part of the theme issue ‘The ecology and evolution of synchronized seed production in plants’.

The ecology and evolution of synchronized reproduction in long-lived plants

Populations of many long-lived plants exhibit spatially synchronized seed production that varies extensively over time, so that seed production in some years is much higher than on average, while in others, it is much lower or absent. This phenomenon termed masting or mast seeding has important consequences for plant reproductive success, ecosystem dynamics and plant–human interactions. Inspired by recent advances in the field, this special issue presents a series of articles that advance the current understanding of the ecology and evolution of masting. To provide a broad overview, we reflect on the state-of-the-art of masting research in terms of underlying proximate mechanisms, ontogeny, adaptations, phylogeny and applications to conservation. While the mechanistic drivers and fitness consequences of masting have received most attention, the evolutionary history, ontogenetic trajectory and applications to plant–human interactions are poorly understood. With increased availability of long-term datasets across broader geographical and taxonomic scales, as well as advances in molecular approaches, we expect that many mysteries of masting will be solved soon. The increased understanding of this global phenomenon will provide the foundation for predictive modelling of seed crops, which will improve our ability to manage forests and agricultural fruit and nut crops in the Anthropocene. This article is part of the theme issue ‘The ecology and evolution of synchronized seed production in plants’.

A brief history of masting research

Although it has long been recognized that seed production by many forest trees varies greatly from year to year, masting (along with ‘mast fruiting’, ‘mast seeding’ and ‘masting behaviour’) as a concept referring to such variability is a relatively recent development. Here, I provide a brief history of masting research, highlighting some of the early contributions by foresters, zoologists and others that paved the way for the burgeoning number of studies currently being conducted by researchers around the world. Of particular current interest is work attempting to understand the proximate mechanisms, evolutionary drivers and community effects of this important ecological phenomenon as well as the ways that climate change may influence masting behaviour in the future. This article is part of the theme issue ‘The ecology and evolution of synchronized seed production in plants'.