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The allocation of resources to plant reproduction or defence at the expense of other fitness traits has been a central component of plant life history theory.

Assessing potential allocation trade-offs among different plant functions is challenging for long-lived plants given the potential for changes in allocation over time.

Despite this challenge, masting species are likely the ideal model for studies of allocation trade-offs given the large, pulsed investment of resources required for reproduction.

Our study focused on a widespread mast-seeding conifer using long-term measures of tree growth alongside cone and resin duct production—traits that are conserved on the surface of limbs or in annual growth—to measure potential trade-offs between these functions across individual, population and landscape scales.

We found evidence for trade-offs among reproduction and defence within individuals, such that trees allocated less resources to defences during mast years.

However, we found no evidence of a growth—reproduction trade-off across all scales, and growth and defence were positively associated at all scales in our study.

We hypothesize that a greater demand for carbohydrates and nutrients in reproduction necessitates a lower allocation to resin duct and terpene production during mast years, while continued allocation to growth would support continued resource allocation and transport.

A key next step for understanding these trade-offs is to evaluate the physiological mechanisms underpinning changing resource allocation between reproductive and defensive pathways within individuals.

Mooney et al. The following additional information is available in the online version of this article—. Figure S1. Relationship between cone production and radial growth left panel and shoot growth right panel.

DEB , D. Breshears DEB and N. Cobb DEB We are grateful for H. Obermueller, A. Shea and L. Hood who provided us with her protocol and accompanying python script to quantify resin duct size, total area and density.

We also thank D. Breshears and N. Cobb, who provided helpful feedback on an earlier draft of this manuscript.

Weevil resistance of progeny derived from putatively resistant and susceptible interior spruce parents.

Forest Ecology and Management : — Google Scholar. Defence reactions of Norway spruce against bark beetles and the associated fungus Ceratocystis polonica in secondary pure and mixed species stands.

Forest Ecology and Management : 73 — Fitting linear mixed-effects models using lme4. Journal of Statistical Software 67 : 1 — Allocating resources to reproduction and defense.

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Forest Ecology and Management 49 : — Breheny P , Burchett W. R package version 2. Regional vegetation die-off in response to global-change-type drought.

To grow or to seed: ecotypic variation in reproductive allocation and cone production by young female Aleppo pine Pinus halepensis , Pinaceae.

American Journal of Botany 95 : — Increased moth herbivory associated with environmental stress of pinyon pine at local and regional levels.

Oecologia : — Long-term sexual allocation in herbivore resistant and susceptible pinyon pine Pinus edulis. Oecologia : 78 — Coley PD.

Effects of plant growth rate and leaf lifetime on the amount and type of anti-herbivore defense. Oecologia 74 : — Resource availability and plant antiherbivore defense.

Science : — Influence of female cones on the vegetative growth of Pinus contorta trees. Tree Physiology 6 : — R package version 1.

Google Preview. Relation between cone production and diameter increment of Douglas fir Pseudotsuga memiesii Mirb. Franco , grand fir Abies grandis Dougl.

Canadian Journal of Botany 43 : — ArcGIS desktop: release Influence of region of provenance and climate factors on wood anatomical traits of Pinus nigra Arn.

European Journal of Forest Research : — To grow or defend? Pine seedlings grow less but induce more defences when a key resource is limited.

Tree Physiology 35 : — Resin duct characteristics associated with tree resistance to bark beetles across lodgepole and limber pines.

Ecological Applications 19 : — Forcella F Estimating pinyon cone production in New Mexico and western Oklahoma.

The Journal of Wildlife Management 45 : — Fox J , Weisberg S. Thousand Oaks, CA : Sage. Anatomical and chemical defenses of conifer bark against bark beetles and other pests.

The New Phytologist : — Fritts HC. Tree rings and climate , 1st edn. Cambridge, MA : Academic Press.

Google Earth Engine: planetary-scale geospatial analysis for everyone. Remote Sensing of Environment : 18 — Gross HL.

Crown deterioration and reduced growth associated with excessive seed production by birch. Canadian Journal of Botany 50 : — Costs of defense and their effects on plant productivity.

In: Givnish TJ , ed. On the economy of plant form and function. Gurevitch, J. The carbon—nutrient balance hypothesis: its rise and fall. Ecology Letters 4 : 86 — Genetic parameters of growth and wood quality traits in Picea abies.

Scandinavian Journal of Forest Research 19 : 14 — The dilemma of plants: to grow or defend. The Quarterly Review of Biology 67 : — Hood S , Sala A.

Ponderosa pine resin defenses and growth: metrics matter. Importance of resin ducts in reducing ponderosa pine mortality from bark beetle attack.

Mechanisms to mitigate the trade-off between growth and defense. The Plant Cell 29 : — Keeling CI , Bohlmann J.

Genes, enzymes and chemicals of terpenoid diversity in the constitutive and induced defence of conifers against insects and pathogens. Resistance of Sitka spruce Picea sitchensis Bong.

Forestry 84 : 83 — Kläy M. Are defensive structures good predictors of tree mortality under drought and insect pressure?

Klutsch JG , Erbilgin N. Dwarf mistletoe infection in jack pine alters growth-defense relationships. Tree Physiology 38 : — Scale of mast-seeding and tree-ring growth.

Nature : — Patterns of annual seed production by Northern hemisphere trees: a global perspective. The American Naturalist : 59 — Koricheva J.

Ecology 83 : — Genetic variation in space and time in a population of ponderosa pine. Heredity 46 : — Environmental effects on constitutive and inducible resin defences of Pinus taeda.

Ecology Letters 3 : — Loomis WE. Growth-differentiation balance vs carbohydrate-nitrogen ratio. Proceedings of the American Society for Horticultural Science 29 : — Lorio PL.

Growth-differentiation balance: a basis for understanding southern pin beetle-tree interactions. Forest Ecology and Management 14 : — Variation in wood anatomical traits of Pinus sylvestris L.

Trees Structure and Function 24 : — The New Phytologist : 91 — Monks A , Kelly D. Austral Ecology 31 : — Masting in ponderosa pine: comparisons of pollen and seed over space and time.

Differential allocation of constitutive and induced chemical defenses in pine tree juveniles: a test of the optimal defense theory.

PLoS One 7 : e Differentiation of persistent anatomical defensive structures is costly and determined by nutrient availability and genetic growth-defence constraints.

Moustakas A , Evans MR. Effects of growth rate, size, and light availability on tree survival across life stages: a demographic analysis accounting for missing values and small sample sizes.

BMC Ecology 15 : 6. Neale DB , Kremer A. Forest tree genomics: growing resources and applications. Nature Reviews Genetics 12 : — Niinemets Ü.

A review of light interception in plant stands from leaf to canopy in different plant functional types and in species with varying shade tolerance.

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Canadian Journal of Botany 47 : — Ecosphere 9 : e Mechanisms of mast seeding: resources, weather, cues, and selection. R package version 3.

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This receives data inputs from the pilot, signals from sensors, and EFIS format selections made by the pilot. The symbol generator can go by other names, such as display processing computer, display electronics unit, etc.

The symbol generator does more than generate symbols. It has at the least monitoring facilities, a graphics generator and a display driver. The required computations are performed, and the graphics generator and display driver produce the inputs to the display units.

Like personal computers, flight instrument systems need power-on-self-test facilities and continuous self-monitoring.

Flight instrument systems, however, need additional monitoring capabilities:. Traditional electromechanical displays are equipped with synchro mechanisms that transmit the pitch, roll, and heading shown on the captain and first officer's instruments to an instrument comparator.

The comparator warns of excessive differences between the Captain and First Officer displays. Even a fault as far downstream [3] as a jam in, say, the roll mechanism of an ADI triggers a comparator warning.

The instrument comparator thus provides both comparator monitoring and display monitoring. With EFIS, the comparator function is simple: Is roll data bank angle from sensor 1 the same as roll data from sensor 2?

Comparison monitors give warnings for airspeed, pitch, roll, and altitude indications. More advanced EFIS systems have more comparator monitors.

In this technique, each symbol generator contains two display monitoring channels. One channel, the internal, samples the output from its own symbol generator to the display unit and computes, for example, what roll attitude should produce that indication.

Any difference has probably been introduced by faulty processing, and triggers a warning on the relevant display.

The external monitoring channel carries out the same check on the symbol generator on the other side of the flight deck: the Captain's symbol generator checks the First Officer's, the First Officer's checks the Captain's.

Whichever symbol generator detects a fault, puts up a warning on its own display. The external monitoring channel also checks sensor inputs to the symbol generator for reasonableness.

A spurious input, such as a radio height greater than the radio altimeter's maximum, results in a warning. At various stages of a flight, a pilot needs different combinations of data.

Ideally, the avionics only show the data in use—but an electromechanical instrument must be in view all the time.

Under normal conditions, an EFIS might not display some indications, e. Only when some parameter exceeds its limits does the system display the reading.

In the case of an input failure, an electromechanical instrument adds yet another indicator—typically, a bar drops across the erroneous data.

EFIS, on the other hand, removes invalid data from the display and substitutes an appropriate warning. A de-clutter mode activates automatically when circumstances require the pilot's attention for a specific item.

For example, if the aircraft pitches up or down beyond a specified limit—usually 30 to 60 degrees—the attitude indicator de-clutters other items from sight until the pilot brings the pitch to an acceptable level.

This helps the pilot focus on the most important tasks. Traditional instruments have long used color, but lack the ability to change a color to indicate some change in condition.

The electronic display technology of EFIS has no such restriction and uses color widely. For example, as an aircraft approaches the glide slope, a blue caption can indicate glide slope is armed, and capture might change the color to green.

Typical EFIS systems color code the navigation needles to reflect the type of navigation. Magenta needles indicate GPS navigation.

EFIS provides versatility by avoiding some physical limitations of traditional instruments. A pilot can switch the same display that shows a course deviation indicator to show the planned track provided by an area navigation or flight management system.

Pilots can choose to superimpose the weather radar picture on the displayed route. The flexibility afforded by software modifications minimises the costs of responding to new aircraft regulations and equipment.

Software updates can update an EFIS system to extend its capabilities. Updates introduced in the s included the ground proximity warning system and traffic collision avoidance system.

A degree of redundancy is available even with the simple two-screen EFIS installation. Should the PFD fail, transfer switching repositions its vital information to the screen normally occupied by the navigation display.

Recent advances in computing power and reductions in the cost of liquid-crystal displays and navigational sensors such as GPS and attitude and heading reference system have brought EFIS to general aviation aircraft.

The low cost is possible because of steep drops in the price of sensors and displays, and equipment for experimental aircraft doesn't require expensive Federal Aviation Administration certification.

This latter point restricts their use to experimental aircraft and certain other aircraft categories, depending on local regulations.

Uncertified EFIS systems are also found in Light-sport aircraft , including factory built, microlight, and ultralight aircraft.

These systems can be fitted to certified aircraft in some cases as secondary or backup systems depending on local aviation rules.

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