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Predictor variables were always z -scored prior to analyses and thus standardized regression coefficients are reported.
We assessed trade-offs among reproduction, growth and defence across trees using a similar linear mixed effect modelling approach as our analyses above.
Specifically, we performed three separate analyses to model seed cone production and our two defence metrics. The predictor variables in each model included the two growth metrics, the two defence metrics cone production model only and cone production defence models only.
Basal area of each tree was also included as a predictor variable in each model to account for potential differences in tree size influencing cone production or our defence metrics.
For these analyses, we quantified cone production, growth and defence metrics for each tree by averaging across all years. Cone production was calculated as the mean number of cones produced per year and thus modelled as a continuous variable rather than a binary variable.
All cone production and defence models were modelled with a Gaussian distribution and an intercept for site was included as a random effect.
We performed correlation analyses to assess for trade-offs between reproduction, growth and defence across sites. Cone production, growth and defence metrics for each site were quantified by averaging the data across all trees within a site.
We found no evidence that climate conditions associated with forest drought stress e. Histograms show how the frequency secondary y -axis of years with above-average cone production top histogram and below-average cone production bottom histogram varies by total resin duct area.
The highest histogram bin includes all data with a resin duct area greater than 0. Each data point is 1 year of data nested within each tree and nested within each site and thus shows relationships at the annual level.
Each data point is a tree and data points of the same colour are trees all within the same site, with the colours matching the sites in Fig.
There was one outlier in panel A see top right of figure and the grey lines show the modelled relationship with the outlier included in the analyses, whereas the black lines in panel A show the modelled relationship with the outlier excluded.
We found no evidence for trade-offs between tree growth, reproduction and defence across our eight sites. 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. Weevil resistance of progeny derived from putatively resistant and susceptible interior spruce parents.
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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. From Wikipedia, the free encyclopedia.
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