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These results suggest that sites with greater growth rates on average were generally more defended. Relationship between mean site tree defence metrics total resin duct area and mean resin duct size and mean site growth metrics radial growth and primary shoot growth.
Each data point is a site, calculated as the mean value among all trees sampled within that site, and light grey brackets show the standard error.
Due to their long lifespans, trees are subject to a dynamic array of attacks from natural enemies as well as changing environmental conditions that alter resource availability.
How trees balance the production of growth, defence and reproduction by allocating among these costly functions is thus central to our understanding of forest ecology.
At the same time, resin duct defences have been demonstrated to have clear importance for pine survival in the face of bark beetle attacks Kane and Kolb ; Kläy ; Ferrenberg et al.
Nevertheless, our results indicate that seed production is favoured over defences, despite a risk of exposure to natural enemies.
This apparently risky trade-off is consistent with the resource-switching hypothesis Pearse et al. Alternatively, this trade-off may be necessary in order to develop large enough seed crops to satiate predators or increased pollination efficiency i.
The resource scarcity in these semi-arid ecosystems may make it particularly important to allocate resources to pulsed, large reproductive efforts during favourable climatic or resource conditions, regardless of exposure to natural enemies.
The negative association between defence investment and cone production suggests that the resources needed to produce these are both limited and linked, unlike xylem growth.
Sala et al. Our results, however, join those from a majority of previous studies that have examined resin duct characteristics of pines and revealed positive growth—defence relationships whereby more growth led to an increase in resin duct production, size or total area within the xylem Kane and Kolb ; Ferrenberg et al.
The association between total resin duct area and radial xylem growth in our study and others may be partly tautological as there is greater space for resin ducts, although notably we also found a similar positive relationship between primary shoot growth and total resin duct area.
This positive relationship of growth and defence indicates that factors which promote more growth, such as greater nutrient or water availability, also lead to more resin duct defence production.
This finding is also supported by a meta-analysis which found that terpenoid-based defences tend to have a positive relationship to growth and only exhibit negative relationships when resources are highly abundant Koricheva —a result predicted by the now-defunct carbon—nutrient balance hypothesis Hamilton et al.
Pine defences are subject to adaptive and directional selection, as both resin duct characteristics and overall production of resin are genetically controlled and at least a moderate amount of the observed phenotypic variation has been shown to be heritable in congeners of P.
In previous studies, resin duct traits have been shown to vary across pine populations Martin et al. Additionally, evidence from Norway and Sitka spruce also indicates that resin duct characteristics can significantly vary among genotypes from the same family group Hannrup et al.
Despite this earlier work, we did not find trade-offs in resource allocation among trees or among populations, only across years.
This study provides strong support for trade-offs between defence and reproduction among individuals yet failed to detect trade-offs across individuals or between growth and defence or reproduction.
The lack of these other trade-offs may be partially due to the limitations of our study design. Most importantly, this study was observational and as a result we were unable to control differences in resource availability across trees or populations.
The positive associations between tree growth and defence may thus be due to microsites with greater resource availability, allowing a given tree to allocate more resources to both growth and defence.
Another key limitation is the cone abscission scar method used to reconstruct historic cone production. Whereas this method is highly effective at reconstructing past cone production Redmond et al.
As a result, there may have been years that trees stopped allocating resources to seed production, leading to noise in our model and thus reducing our ability to detect trade-offs at the ultimate level of embryos.
In addition, we were unable to sample trees that had highly damaged branches and thus no available markers of cone production, which may have been exceptionally poor or high producers of resin ducts or cone production.
Our metrics of defence allocation were also limited to the amount of defence structures produced and did not include variation in monoterpene production, composition and volatile emissions from resin that all contribute to a trees ability to defend against insect infestations Keeling and Bohlmann These limitations underscore the importance of continued research on trade-offs in resource allocations given the challenges of observational studies, especially those reconstructing past investment in growth, reproduction and defence.
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.
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Resin yield in Pinus pinaster is related to tree dendrometry, stand density and tapping-induced systemic changes in xylem anatomy. EFIS provides pilots with controls that select display range and mode for example, map or compass rose and enter data such as selected heading.
Where other equipment uses pilot inputs, data buses broadcast the pilot's selections so that the pilot need only enter the selection once.
For example, the pilot selects the desired level-off altitude on a control unit. The EFIS repeats this selected altitude on the PFD, and by comparing it with the actual altitude from the air data computer generates an altitude error display.
This same altitude selection is used by the automatic flight control system to level off, and by the altitude alerting system to provide appropriate warnings.
The EFIS visual display is produced by the symbol generator. 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  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.