Freezing And Mechanical Failure of A Habitat-forming Kelp in The Rocky Intertidal Zone.

Kelp and other habitat-forming seaweeds in the intertidal zone are exposed to a suite of environmental factors, including temperature and hydrodynamic forces, that can influence their growth, survival, and ecological function. Relatively little is known about the interactive effect of temperature and hydrodynamic forces on kelp, especially the effect of cold stress on biomechanical resistance to hydrodynamic forces. We used the intertidal kelp Egregia menziesii to investigate how freezing in air during a low tide changes the kelp's resistance to breaking from hydrodynamic forces. We conducted a laboratory experiment to test how short-term freezing, mimicking a brief low tide freezing event, affected the kelp's mechanical properties. We also characterized daily minimum winter temperatures in an intertidal E. menziesii population on San Juan Island, WA near the center of the species' geographic range. In the laboratory, acute freezing events decreased the strength and toughness of kelp tissue by 8 to 20% (change in medians). During low tides in the field, we documented sub-zero temperatures, snow, and low canopy cover (compared to summer surveys). These results suggest that freezing can contribute to frond breakage and decreased canopy cover in intertidal kelp. Further work is needed to understand whether freezing and the biomechanical performance in cold temperatures influence the fitness and ecological function of kelp, and whether this will change as winter conditions, such as freezing events and storms, change in frequency and intensity.

Flying, nectar-loaded honey bees conserve water and improve heat tolerance by reducing wingbeat frequency and metabolic heat production.

Heat waves are becoming increasingly common due to climate change, making it crucial to identify and understand the capacities for insect pollinators, such as honey bees, to avoid overheating. We examined the effects of hot, dry air temperatures on the physiological and behavioral mechanisms that honey bees use to fly when carrying nectar loads, to assess how foraging is limited by overheating or desiccation. We found that flight muscle temperatures increased linearly with load mass at air temperatures of 20 or 30 °C, but, remarkably, there was no change with increasing nectar loads at an air temperature of 40 °C. Flying, nectar-loaded bees were able to avoid overheating at 40 °C by reducing their flight metabolic rates and increasing evaporative cooling. At high body temperatures, bees apparently increase flight efficiency by lowering their wingbeat frequency and increasing stroke amplitude to compensate, reducing the need for evaporative cooling. However, even with reductions in metabolic heat production, desiccation likely limits foraging at temperatures well below bees' critical thermal maxima in hot, dry conditions.

Close encounters of three kinds: impacts of leg, wing and body collisions on flight performance in carpenter bees.

Flying insects often forage among cluttered vegetation that forms a series of obstacles in their flight path. Recent studies have focused on behaviors needed to navigate clutter while avoiding all physical contact and, as a result, we know little about flight behaviors that do involve encounters with obstacles. Here, we challenged carpenter bees (Xylocopa varipuncta) to fly through narrow gaps in an obstacle course to determine the kinds of obstacle encounters they experience, as well as the consequences for flight performance. We observed three kinds of encounters: leg, body and wing collisions. Wing collisions occurred most frequently (in about 40% of flights, up to 25 times per flight) but these had little effect on flight speed or body orientation. In contrast, body and leg collisions, which each occurred in about 20% of flights (1-2 times per flight), resulted in decreased flight speeds and increased rates of body rotation (yaw). Wing and body collisions, but not leg collisions, were more likely to occur in wind versus still air. Thus, physical encounters with obstacles may be a frequent occurrence for insects flying in some environments, and the immediate effects of these encounters on flight performance depend on the body part involved.

Ecological biomechanics of damage to macroalgae.

Macroalgae provide food and habitat to a diversity of organisms in marine systems, so structural damage and breakage of thallus tissue can have important ecological consequences for the composition and dynamics of marine communities. Common sources of macroalgal damage include breakage by hydrodynamic forces imposed by ambient water currents and waves, tissue consumption by herbivores, and injuries due to epibionts. Many macroalgal species have biomechanical designs that minimize damage by these sources, such as flexibly reconfiguring into streamlined shapes in flow, having either strong or extensible tissues that are tough, and having chemical and morphological defenses against herbivores and epibionts. If damage occurs, some macroalgae have tissue properties that prevent cracks from propagating or that facilitate tissue breakage in certain places, allowing the remainder of the thallus to survive. In contrast to these mechanisms of damage control, some macroalgae use breakage to aid dispersal, while others simply complete their reproduction prior to seasonally-predictable periods of damage (e.g., storm seasons). Once damage occurs, macroalgae have a variety of biomechanical responses, including increasing tissue strength, thickening support structures, or altering thallus shape. Thus, macroalgae have myriad biomechanical strategies for preventing, controlling, and responding to structural damage that can occur throughout their lives.

An evaluation of common methods for comparing the scaling of vertical force production in flying insects.

Maximum vertical force production (Fvert) is an integral measure of flight performance that generally scales with size. Numerous methods of measuring Fvert and body size are accessible to entomologists, but we do not know whether method selection affects inter- and intraspecific comparisons of Fvert-size scaling. We compared two common techniques for measuring Fvert in bumblebees (Bombus impatiens) and mason bees (Osmia lignaria), and examined Fvert scaling using five size metrics. Fvert results were similar with incremental or asymptotic load-lifting, but scaling analyses were sensitive to the size metric used. Analyses based on some size metrics indicated similar scaling exponents and coefficients between species, whereas other metrics indicated coefficients that differed by up to 18%. Furthermore, Fvert showed isometry with body lengths and fed and starved masses, but negative allometry with dry mass. We conclude that Fvert can be measured using either incremental or asymptotic loading but choosing a size metric for scaling studies requires careful consideration.

Wind and route choice affect performance of bees flying above versus within a cluttered obstacle field.

Bees flying through natural landscapes frequently encounter physical challenges, such as wind and cluttered vegetation, but the influence of these factors on flight performance remains unknown. We analyzed 548 videos of wild-caught honeybees (Apis mellifera) flying through an enclosure containing a field of vertical obstacles that bees could choose to fly within (through open corridors, without maneuvering) or above. We varied obstacle field height and wind condition (still, headwinds or tailwinds), and examined how these factors affected bees' flight altitude, ground speed, and side-to-side casting motions (lateral excursions). When obstacle fields were short, bees flew at altitudes near the midpoint between the tunnel floor and ceiling. When obstacle fields approached or exceeded this midpoint, bees tended to increase their altitude, but they did not always avoid flying through obstacles, despite having the freedom to do so. Bees that flew above the obstacles exhibited 40% faster ground speeds and 36% larger lateral excursions than bees that flew within the obstacle fields. Wind did not affect flight altitude, but bees flew 12-19% faster in tailwinds, and their lateral excursions were 19% larger when flying in headwinds or tailwinds, as compared to still air. Our results show that bees flying through complex environments display flexibility in their route choices (i.e., flying above obstacles in some trials and through them in others), which affects their overall flight performance. Similar choices in natural landscapes could have broad implications for foraging efficiency, pollination, and mortality in wild bees.

A push for inclusive data collection in STEM organizations.

Professional societies could better survey, and thus better serve, underrepresented groups.

Flow, form, and force: methods and frameworks for field studies of macroalgal biomechanics.

Macroalgae are ecologically important organisms that often inhabit locations with physically challenging water motion. The biomechanical traits that permit their survival in these conditions have been of interest to biologists and engineers alike, but logistical and technical challenges of conducting investigations in macroalgal habitats have often prevented optimal study of these traits. Here, we review field methods for quantifying three major components of macroalgal biomechanics in moving water: fluid flow, macroalgal form, and hydrodynamic force. The implementation of some methodologies is limited due to the current state and accessibility of technology, but many of these limitations can be remedied by custom-built devices, borrowing techniques from other systems, or shifting lab-based approaches to the field. We also describe several frameworks for integrating flow, form, and force data that can facilitate comparisons of macroalgal biomechanics in field settings with predictions from theory and lab-based experiments, or comparisons between flow conditions, habitats, and species. These methods and frameworks, when used on scales that are relevant to the examined processes, can reveal mechanistic information about the functional traits that permit macroalgae to withstand physically challenging water motion in their habitats, using the actual fluid flows, macroalgal forms, and physical forces that occur in nature.

Kelp Morphology and Herbivory Are Maintained Across Latitude Despite Geographic Shift in Kelp-Wounding Herbivores.

AbstractHerbivores can drastically alter the morphology of macroalgae by directly consuming tissue and by inflicting structural wounds. Wounds can result in large amounts of tissue breaking away from macroalgae, amplifying the damage initially caused by herbivores. Herbivores that commonly wound macroalgae often occur over only a portion of a macroalga's lifespan or geographic range. However, we know little about the influence of these periodic or regional occurrences of herbivores on the large-scale seasonal and geographical patterns of macroalgal morphology. We used the intertidal kelp Egregia menziesii to investigate how the kelp's morphology and the prevalence of two prominent kelp-wounding herbivores (limpets and amphipods) changed over two seasons (spring and summer) and over the northern extent of the kelp's geographic range (six sites from central California to northern Washington). Wounds from limpets and amphipods often result in the kelp's fronds being pruned (intercalary meristem broken away), so we quantified kelp size (combined length of all fronds) and pruning (proportion of broken fronds). We found similar results in each season: herbivores were most likely to occur on large, pruned kelp regardless of site; and limpets were the dominant herbivore at southern sites, while amphipods were dominant at northern sites. Despite the geographic shift in the dominant herbivore, kelp had similar levels of total herbivore prevalence (limpets and/or amphipods) and similar morphologies across sites. Our results suggest that large-scale geographic similarities in macroalgal wounding, despite regional variation in the herbivore community, can maintain similar macroalgal morphologies over large geographic areas.

Age affects the strain-rate dependence of mechanical properties of kelp tissues.

PREMISE: The resistance of macroalgae to hydrodynamic forces imposed by ambient water motion depends in part on the mechanical properties of their tissues. In wave-swept habitats, tissues are stretched (strained) at different rates as hydrodynamic forces change. Previous studies of mechanical properties of macroalgal tissues have used either a single strain rate or a small range of strain rates. Therefore, our knowledge of the mechanical properties of macroalgae is limited to a narrow fraction of the strain rates that can occur in nature. In addition, although mechanical properties of macroalgal tissues change with age, the effect of age on the strain-rate dependence of their mechanical behavior has not been documented. METHODS: Using the kelp Egregia menziesii, we measured how high strain rate (simulating wave impingement) and low strain rate (simulating wave surge) affected mechanical properties of frond tissues of various ages. RESULTS: Stiffness of tissues of all ages increased with strain rate, whereas extensibility was unaffected. Strength and toughness increased with strain rate for young tissue but were unaffected by strain rate for old tissue. CONCLUSIONS: Young tissue is weaker than old tissue and, therefore, the most susceptible to breakage from hydrodynamic forces. The increased strength of young tissue at high strain rates can help the frond resist breaking when pulled rapidly during wave impingement, when hydrodynamic forces are largest. Because breakage of young tissue can remove a frond's meristem and negatively impact the survival of the whole kelp, strain-rate dependence of the young tissue's strength can enhance kelp's survival.

Wind and obstacle motion affect honeybee flight strategies in cluttered environments.

Bees often forage in habitats with cluttered vegetation and unpredictable winds. Navigating obstacles in wind presents a challenge that may be exacerbated by wind-induced motions of vegetation. Although wind-blown vegetation is common in natural habitats, we know little about how the strategies of bees for flying through clutter are affected by obstacle motion and wind. We filmed honeybees Apis mellifera flying through obstacles in a flight tunnel with still air, headwinds or tailwinds. We tested how their ground speeds and centering behavior (trajectory relative to the midline between obstacles) changed when obstacles were moving versus stationary, and how their approach strategies affected flight outcome (successful transit versus collision). We found that obstacle motion affects ground speed: bees flew slower when approaching moving versus stationary obstacles in still air but tended to fly faster when approaching moving obstacles in headwinds or tailwinds. Bees in still air reduced their chances of colliding with obstacles (whether moving or stationary) by reducing ground speed, whereas flight outcomes in wind were not associated with ground speed, but rather with improvement in centering behavior during the approach. We hypothesize that in challenging flight situations (e.g. navigating moving obstacles in wind), bees may speed up to reduce the number of wing collisions that occur if they pass too close to an obstacle. Our results show that wind and obstacle motion can interact to affect flight strategies in unexpected ways, suggesting that wind-blown vegetation may have important effects on foraging behaviors and flight performance of bees in natural habitats.

Physiological responses to heat stress in an invasive mussel Mytilus galloprovincialis depend on tidal habitat.

Mussels are ecologically important organisms that can survive in subtidal and intertidal zones where they experience thermal stress. We know little about how mussels from different tidal habitats respond to thermal stress. We used the mussel Mytilus galloprovincialis from separate subtidal and intertidal populations to test whether heart rate and indicators of potential aerobic (citrate synthase activity) and anaerobic (cytosolic malate dehydrogenase activity) metabolic capacity are affected by increased temperatures while exposed to air or submerged in water. Subtidal mussels were affected by warming when submerged in water (decreased heart rate) but showed no effect in air. In contrast, intertidal mussels were affected by exposure to air (increased anaerobic capacity) but not by warming. Overall, physiological responses of mussels to thermal stress were dependent on their tidal habitat. These results highlight the importance of considering the natural habitat of mussels when assessing their responses to environmental challenges.

Turbinaria ornata (Phaeophyceae) varies size and strength to maintain environmental safety factor across flow regimes.

Water motion in coastal areas can produce hydrodynamic forces that damage or dislodge benthic macroalgae if the tissues of macroalgae are not sufficiently strong. Some macroalgae vary their morphology and strength in response to ambient water motion, but little is known of how morphology and strength of macroalgae change relative to one another across flow regimes. Here, we use Turbinaria ornata, an ecologically important macroalga, to study how both the morphology and strength of macroalgae vary with ambient water motion. Typically, T. ornata exhibits weakening of its stipe when sexually mature, leading to breakage from the substratum and dispersal, which is beneficial for reproduction. Across three flow regimes, adult T. ornata increased its size but decreased its strength as water motion decreased. However, the strength of T. ornata relative to the maximum hydrodynamic forces it is expected to encounter (the environmental safety factor) did not differ between flow regimes. Our results showed that T. ornata can conform to its local flow habitat by varying both size and strength, similar to other macroalgae. Varying multiple traits between flow regimes suggested that T. ornata is capable of surviving a wide range of flow conditions, which may permit more control over the timing of its weakening, breakage from the substratum, and dispersal, even with future increases in flow velocities (e.g., large waves from storms) that are expected to occur frequently with climate change.

Functional responses of intertidal bivalves to repeated sub-lethal, physical disturbances.

In coastal habitats, physical disturbances of benthic organisms can be caused by natural events like wave-born objects and human activity like trampling, and these disturbances can be sub-lethal (e.g., resulting in the organism's displacement). We know little of how sessile organisms respond to physical disturbances such as displacements. Using Mytilaster minimus, a mussel that is native to the Mediterranean Sea, we tested how byssus production and oxygen uptake rates changed in response to different frequencies of disturbance events (10-60 events h-1). Mussels increased oxygen uptake rates but not byssus production with increasing disturbance frequencies (50-60 events h-1). Our results show that sub-lethal, physical disturbances can cause increased physiological rates in mussels if disturbances repeat rapidly. Therefore, sub-lethal, physical disturbances can have negative consequences for benthic organisms even if they do not cause immediate damage or mortality.

Mechanical properties of the wave-swept kelp Egregia menziesii change with season, growth rate and herbivore wounds.

The resistance of macroalgae to damage by hydrodynamic forces depends on the mechanical properties of their tissues. Although factors such as water-flow environment, algal growth rate and damage by herbivores have been shown to influence various material properties of macroalgal tissues, the interplay of these factors as they change seasonally and affect algal mechanical performance has not been worked out. We used the perennial kelp Egregia menziesii to study how the material properties of the rachis supporting a frond changed seasonally over a 2 year period, and how those changes correlated with seasonal patterns of the environment, growth rate and herbivore load. Rachis tissue became stiffer, stronger and less extensible with age (distance from the meristem). Thus, slowly growing rachises were stiffer, stronger and tougher than rapidly growing ones. Growth rates were highest in spring and summer when upwelling and long periods of daylight occurred. Therefore, rachis tissue was most resistant to damage in the winter, when waves were large as a result of seasonal storms. Herbivory was greatest during summer, when rachis growth rates were high. Unlike other macroalgae, E. menziesii did not respond to herbivore damage by increasing rachis tissue strength, but rather by growing in width so that the cross-sectional area of the wounded rachis was increased. The relative timing of environmental factors that affect growth rates (e.g. upwelling supply of nutrients, daylight duration) and of those that can damage macroalgae (e.g. winter storms, summer herbivore outbreaks) can influence the material properties and thus the mechanical performance of macroalgae.

Herbivores alter plant-wind interactions by acting as a point mass on leaves and by removing leaf tissue.

In nature, plants regularly interact with herbivores and with wind. Herbivores can wound and alter the structure of plants, whereas wind can exert aerodynamic forces that cause the plants to flutter or sway. While herbivory has many negative consequences for plants, fluttering in wind can be beneficial for plants by facilitating gas exchange and loss of excess heat. Little is known about how herbivores affect plant motion in wind. We tested how the mass of an herbivore resting on a broad leaf of the tulip tree Liriodendron tulipifera, and the damage caused by herbivores, affected the motion of the leaf in wind. For this, we placed mimics of herbivores on the leaves, varying each herbivore's mass or position, and used high-speed video to measure how the herbivore mimics affected leaf movement and reconfiguration at two wind speeds inside a laboratory wind tunnel. In a similar setup, we tested how naturally occurring herbivore damage on the leaves affected leaf movement and reconfiguration. We found that the mass of an herbivore resting on a leaf can change that leaf's orientation relative to the wind and interfere with the ability of the leaf to reconfigure into a smaller, more streamlined shape. A large herbivore load slowed the leaf's fluttering frequency, while naturally occurring damage from herbivores increased the leaf's fluttering frequency. We conclude that herbivores can alter the physical interactions between wind and plants by two methods: (1) acting as a point mass on the plant while it is feeding and (2) removing tissue from the plant. Altering a plant's interaction with wind can have physical and physiological consequences for the plant. Thus, future studies of plants in nature should consider the effect of herbivory on plant-wind interactions, and vice versa.