
Common Name: Masting – The production of copious quantities of deciduous tree nuts in a single year followed by several years of minimal nut production. The phenomenon involves all nut trees, regardless of species, within a well-defined geographical region.
Scientific Name: Seed masting or Mast Seeding – A slightly more technical term to emphasize that the purpose of masting is to accentuate seed propagation to promote new tree growth.
Potpourri: Mast is a noun of Anglo-Saxon origin (mæst in the original Old English form using the ligature æ) that refers to the accumulation of various kinds of nuts on the forest floor that served as food for farm animals, particularly domesticated hogs. Pannage is a mostly arcane term for the pasturing of animals in the forest to take advantage of the mast, a practice that played a major role in the sociological development of rural life in Europe. As the swine population grew in concert with the human population, the pannage season had to be restricted, traditionally from the feast of Saint Michael (September 29) to the last day of November. Pigs were brought to the New World with the earliest expeditions, notably that of Hernando De Soto in the southeast from 1539 to 1542, the progenitors of the razorback. Hogs became central to the salt pork and fatback cultures of the Appalachian and Ozark Mountains, the oak-chestnut forests providing the mast for their sustenance. [1]
The process by which trees produce mast is called masting. The curious thing about masting is that it is not a continuous process but rather is cyclic. Every three to five years a tree will produce prodigious quantities of nuts; in between the “masts,” it will produce almost none. The span of time between masts varies according to tree species and a host of other ecological and climate factors and can be as long as ten years. It is a matter of common experience that many kinds of trees exhibit this behavior at the same time over a large geographic area. This poses two conundrums: (1) Why do the trees regulate their nut production in a boom or bust manner?; and (2) How do they manage to coordinate the same cycle with other trees over a large area? Individual tree masting is called variability and the coordination among masting trees is called synchrony. [2]
Variability has had two hypothetical explanations: resource responsiveness and economy of scale. The basic precept of resource responsiveness is that an individual tree will respond to the resources at its disposal. In a good year with plenty of rain and sunlight, a tree would have more resources with which to manufacture more nuts, which would subsequently be more likely to propagate in a moist, nutrient rich environment. The primary resources of interest (rain and an adequacy of sunlight) are determined by prevailing weather conditions. Since weather patterns extend over an extended geographic area, resource tracking could also explain synchrony, as all the trees would be subject to the same cycle of resources.
Nature is not that simple, however. The fact is that variations in weather do not correlate with masting; moist and sunny weather does not produce a mast crop any more than dry and overcast weather prevents one. Weather is not cyclic; a wet year is not necessarily followed by a dry year. Masting is much more consistent in periodicity and result, cycles of high nut production occurring on a regular, periodic basis. However, there is one aspect of resource utilization by masting trees that does track with mast cycles, the resources expended by the tree. A significant resource investment (estimated to be about 10 percent of its total nutrients) must be made by a tree to produce the flowers that, when fertilized, produce the seed nuts. What this means is that trees grow slowly during mast years and more rapidly in non-mast years as the resources are shifted from reproduction to growth. This suggests that masting is part of a complex evolutionary behavior pattern that must derive from an ecological stimulus – economy of scale variability.
The term economy of scale refers to the general precept that benefits will be magnified by the scale of the population size. Buying in bulk lots distinguishes wholesale from retail with the former gaining the economy of scale reduced costs of larger quantities. There are two corollaries associated with the economy of scale theory for masting variability – predator satiation and pollination efficiency. In predator satiation, masting is stimulated by a tree’s adaptive strategy for survival in a world of nut-eating predators (notably squirrels). By producing a super-sized nut crop, the predators become satiated so that an adequate number of nuts will survive to succeed for propagation of the tree species.
The seven white oaks in my backyard so overwhelm the squirrel population with acorns in a mast year that they scamper about in confusion with too many nuts to eat or bury. A rough calculation based on extrapolation yields a total over 200,000 acorns. When squirrels cross from the back yard to the front past the side of the house, they are confronted with an equally overwhelming mass of hickory nuts from the hickory trees there, which, of course, mast in synchrony with the oak trees. The predator population is held in check during the non-mast years, when the parsimony of production is reflected in declining populations. In the economy of scale paradigm, each nut in a mast year has a greater probability of escaping predation.

Pollination efficiency is based on the notion that it is more efficient from the resource standpoint for a plant to successfully propagate if there are a large number of sites for germination. This is not true for all plants. Flowering (Angiosperm) plants employ insect pollination, meting out nectar advertised by their attractive flowers to take advantage of male pollen transport to the female ovary of another plant. Chicory is a good example, as only a few flowers open each day and each expires at day’s end. Oak and hickory trees are monecious (male and female flowers are on the same tree) and their pollen is transported from staminate to pistillate flowers by the wind. From the standpoint of reproductive success, it is advantageous for oaks to fill the air with pollen from many trees at the same time, saving up energy during off-years. Fungi are also mostly wind-pollinated and accordingly produce spores in prodigal proportions; a giant puffball is estimated to contain about 7 trillion spores.
Field testing for experimental evidence of predator satiation and pollination efficiency as causative factors for the masting behavior of trees is difficult and the results tenuous. For example, a study of masting trees in a 6-hectare study area estimated pollination efficiency by counting the total number male flowers and the number of nuts produced from1988 to 1993. Testing for predator satiation is even more difficult; one must not only show that predators were satiated but also that the interval between masting events was sufficient to result in a decrease in predator population. The same study utilized the number of nuts that had evidence of insect predation relative to those that were undamaged as a measure of satiation. The year-to-year variance in nuts with evidence of insect predation was used to determine the mast interval. The study concluded that both effects were observable, pollination efficiency having a greater impact than predator satiation. [3]
But the real conundrum is not why trees mast as individuals (variability), but how they coordinate their activities over large areas and across different species (synchrony). It is a matter of direct observation and scientific study that they do. A survey of acorn production was initiated in 1994 to quantify acorn production of blue oaks at 10 different sites at separated by up to 700 kilometers in California. The conclusion, after eleven years of study was that “acorn production extends to pretty much every blue oak, a population of 100 to 200 million individuals.” A more comprehensive literature survey of relevant references on nut production by various trees was organized by W.D. Koenig, a professor at UC Berkeley. A review of 72 sites and 5,000 data points revealed that synchronization of seed production was statistically significant in populations separated by 2500 kilometers. [4] One may conclude that synchrony occurs over long distances and involves almost every tree. So how do they do it?
Three mechanisms are germane to any discussion of synchronization of activities among plants or animals: chemical, reproductive, and environmental. The use of chemicals to transmit signals among individuals is common. However, it is not likely that this is pertinent to the case of masting as chemicals act over much shorter ranges than is observed in masting tree populations. Reproductive synchronization in arboreal terms is called pollen coupling. The concept is that if a tree depends on the pollen from a second tree to produce the fruit nut, then it must be synchronized with it. Implicit in this is that the tree that is providing the pollen must be at some relevant distance away. The effective distance over which pollen is effective in achieving fertilization is of value in forest management; recently completed studies have revealed that pollen is only effective within a range of about 60 meters, hardly on the order of the observed ranges of masting behavior. [5] A second reason that this mechanism is irrelevant here is that both oak and hickory trees are monecious, so the pollen doesn’t need to travel more than a few meters. Since it is not likely that chemical or reproductive effects result in the long-distance synchrony of masting, one must conclude that the only other reasonable choice would be environmental.
The notion that resource responsiveness to the environment causes the masting behavior of individual trees, i.e. variability, was ruled out above based on field observation. The question is how environmental resources that could not cause masting variability would nonetheless be the cause for masting synchrony. It is the difference between weather and climate, the former term referring to the short-term manifestation of the latter. The idea that the environment can cause synchronous fluctuation in population size is not new. It is called the Moran effect after the Australian statistician who showed that the correlation of two populations at different locations was equal to the correlation in their common environmental influence (if they were subject to the same basic parameters). It has been demonstrated empirically in many organisms, from viruses in the body to caribou in Greenland. [6]
It is therefore possible that geographically wide-ranging climate conditions cause trees to mast in synchrony. It is not known at present what aspect of the climate is predominate, if, in fact, it is that simple. There is some evidence that temperature may be a key parameter. The study of the California oaks was correlated with the mean temperature in April over the course of the eleven years of data. April was chosen as the most important month for masting, as it is when the trees produce the male and female flowers that result in nuts that ultimately fall as mast. The spatial synchronization of April temperatures was found to be even more strongly correlated than the masting of the oaks. The rationale is clear: the periodic fluctuations of temperatures (perhaps caused by the cyclic El Nino phenomenon) operate in synchronization with masting over the same geographic area. [7]
Or maybe it is something else altogether. The recent demonstration of the communion of all of the trees in the forest in concert with their mutualistic fungal partners may offer an alternative hypothesis. It is demonstrably true that the stronger trees in a forest send nutrients to the weaker trees in the forest and the trees in the sun send nutrients to the understory trees in the shade in order to keep the forest ecosystem in balance [8], then why wouldn’t these same signal paths send the signal to make more nuts? As the nutrient levels build up in the tree roots and fungal mycelia during high growth years, a crescendo point is reached and each and every tree gets a boost of energy to make it a mast year. This would certainly explain synchrony, and, given the vast dimensions of the “wood-wide web” it would also explain regional geographic expanse. All of the trees in the forest would benefit from an increased likelihood of the growth of saplings that would eventually succeed their parent trees so that the forest as “mega-organism” survived.
A similar and probably related observation is that many fungi will gather their resources for years only to erupt in a single year that results essentially in what might be considered a mast of mushrooms. There are many similarities as fungal spores, like the pollen of trees, benefit from a large cohort; each germinating spore creates a hypha that must find a mate. The fact that oaks have an especially large number of fungi with which they form mycorrhizal associations is quite likely why they are so successful as climax forest tree species in part due to the benefits of masting to longevity in a healthy forest. [9]
Another point to ponder at this juncture is why the masting phenomenon is restricted only to nut trees like oaks. What about the seed-bearing cones of conifers? It is not uncommon to traverse a pine forest, noting the soft tread of pine needles and a profusion of cones. The pine needles are indicative of a little noted characteristic of evergreens. They lose their needles just as deciduous trees shed their leaves. They just don’t do it all at once but take about four years to recycle all needles (longer for firs and spruces). Cone masting is more of a challenge, however, because it takes three years to produce a cone. Year one requires moisture and sunlight to prime the cone by accumulating resources, year two must be dry to enhance pollination, and year three must be wet and sunny for cone growth. While cone trees “mast” based on field observation experience, there does not appear to be any synchrony between conifers masting cones and deciduous trees masting nuts. This has led to the hypothesis that cone bearing trees produce copious seeds when environmental conditions favor germination as opposed to the more social sharing behavior of nut trees. [10]

Over the last twenty-five years, research to better understand tree masting and its effects of forest health has continued, expanding across the globe from its mostly North America and Europe roots and to involve specialists from fields other than botany. Of note is confirmation of the wide geographic range of masting synchrony that has been shown to be of intercontinental scale, extending to over thousands of kilometers. This suggests that fluctuating weather patterns driven by some environmental factor (the 11-year sunspot cycle has been proposed) must be key to synchrony as nothing else could be so widespread. One of the more influential studies concluded that masting was the fundamental driver of animal behavior: “variable acorn crop size drives a chain reaction linking deer populations, ticks, and Lyme disease along with mouse populations, ground-nesting birds, and gypsy moths”.
The fundamental debate is now focused on the relative importance of resource matching (that masting depends on some limited resource), and economy of scale (that masting depends on genetic evolutionary trends of the tree species). A resource budget model has been proposed that unites the two with advantages of evolving to produce mountains of acorns weighed against the resources needed for synchronous reproduction. [11] The bottom line is that there is not (yet) an accepted comprehensive explanation for masting, despite its importance to the health of forests and the animals and fungi that live in them. It may well be that we are just at the threshold of understanding the real, complex nature of forests.
References:
1. DeVoto, B. The Course of Empire, The Easton Press, Norwalk, Connecticut, 1988. pp 23-31.
2. http://uslancaster.sc.edu/faculty/scarlett/acrnsmry.htm
3. Shibata, M. et al “Causes and Consequences of Mast Seed Production of Four Co-occurring Carpinus species in Japan” Ecology, January 1998, pp 9 – 12 This paper documents a thorough field test of masting hypotheses.
4. Koenig, D. and Knops, J. “The Mystery of Masting in Trees” American Scientist Volume 93 July-August 2005. Pp 340-349.
5. Knapp, E. et al “Pollen-limited reproduction in blue oak: Implications for wind pollination for fragmented applications” Oecologia 128 March 2001 pp 48-55.
6. Moran, P. A. P. “The statistical analysis of the Canadian lynx cycle. II. Synchronization and meteorology” Australian Journal of Zoology, June 1953 pp. 291–298.
7. Koenig, D. and Knops, J. Op. cit.
8. Klein, T. et al “Belowground carbon trade among tall trees in a temperate forest.” Science 15 April 2016, Vol. 352, Issue 6283, pp. 342-344.
9. Binion, D. et al Macrofungi Associated with Oaks of Eastern North America, West Virginia University Press, Morgantown, WV, 2008.
10. Lauder, J. “The Science of Masting: Why are there so many acorns (or cones)?” Sierra Streams Institute. 15 October 2024 https://sierrastreamsinstitute.org/2024/10/15/the-science-of-masting-why-are-there-so-many-acorns-or-cones/
11. Koenig D. A Brief History of Masting Research Philosophical Transactions of the Royal Society, 26 March 2021 Volume 376








