9+ How Fish Survive When a Lake Freezes: Winter Life


9+ How Fish Survive When a Lake Freezes: Winter Life

When surface water temperatures drop below freezing, a layer of ice forms on lakes. This ice cover drastically alters the aquatic environment, impacting the life within. The formation of ice affects light penetration, water temperature, and oxygen availability, all critical factors for the survival of aquatic organisms, particularly fish. The extent and duration of ice cover vary depending on geographical location and climatic conditions.

The consequences of lake freezing are significant for both the fish populations and the broader ecosystem. Understanding these effects is crucial for fisheries management and conservation efforts, especially in regions where lakes are subject to prolonged periods of ice cover. Historically, indigenous communities and early settlers relied on understanding ice formation patterns for winter fishing practices. Current research allows a more detailed and nuanced understanding of how these frozen conditions influence fish physiology and behavior.

This article will delve into the specific physiological adaptations fish employ to survive in frigid, ice-covered waters. It will examine the critical role of dissolved oxygen, the impact of temperature stratification, and the behavioral modifications fish undertake to cope with these challenging conditions. Finally, it will discuss the broader ecological implications of winterkill events and the long-term effects of climate change on these delicate aquatic ecosystems.

1. Reduced Metabolism

The freezing of a lake initiates a cascade of environmental changes that compel fish to enter a state of reduced metabolism. As water temperatures decline, poikilothermic (cold-blooded) fish experience a direct and significant slowing of their metabolic rate. This physiological adaptation is critical for survival under ice because it drastically reduces the fish’s energy requirements and, consequently, its demand for oxygen. The availability of dissolved oxygen in ice-covered lakes often diminishes over time due to the cessation of atmospheric exchange and the ongoing decomposition of organic matter. By lowering their metabolic rate, fish can survive for extended periods on limited oxygen resources. For example, species like crucian carp can enter near-suspended animation in oxygen-deprived environments, surviving for months with minimal energy expenditure.

The degree of metabolic reduction varies among species and is influenced by factors such as body size, age, and prior acclimation to cold temperatures. Smaller fish generally exhibit a higher metabolic rate compared to larger individuals, even under cold conditions, making them potentially more vulnerable to oxygen depletion. Understanding the specific metabolic responses of different fish species is vital for fisheries management. For instance, stocking lakes with species that have a lower metabolic demand during winter can increase the likelihood of successful overwintering and reduce the risk of winterkill events.

In summary, reduced metabolism is an essential survival mechanism that allows fish to endure the harsh conditions imposed by lake freezing. It directly mitigates the challenges of low oxygen and limited food availability. While metabolic depression is advantageous, it also renders fish more susceptible to stress and disease. Research into the metabolic adaptations of fish to cold environments contributes valuable insights for conservation efforts and sustainable management of freshwater ecosystems impacted by seasonal ice cover.

2. Oxygen Depletion

Ice cover acts as a barrier, preventing atmospheric oxygen from dissolving into the water. Simultaneously, the decomposition of organic matter, such as decaying leaves and dead algae, continues consuming dissolved oxygen. This combination creates a situation where oxygen levels gradually decline throughout the winter months. The rate of oxygen depletion depends on factors such as the amount of organic matter present, water temperature, and the duration of ice cover. Shallow lakes with abundant organic sediments are particularly prone to severe oxygen depletion. When oxygen concentrations drop below critical levels for sustained respiration, fish experience stress and, ultimately, suffocation. This phenomenon, termed “winterkill,” can result in significant fish mortality events. Different fish species exhibit varying tolerances to low oxygen conditions; for example, trout and salmon are highly sensitive, while carp and bullheads are relatively tolerant.

The extent of oxygen depletion can be mitigated by the presence of snow cover on the ice. Snow reduces light penetration, inhibiting photosynthetic activity by aquatic plants and algae. While photosynthesis contributes oxygen to the water, excessive algal blooms followed by die-offs can exacerbate oxygen depletion as the dead algae decompose. Management strategies aimed at reducing nutrient loading into lakes can help prevent algal blooms and thereby reduce the risk of winterkill. Artificially aerating lakes through mechanical means or by introducing oxygen can also provide localized relief from oxygen depletion, although this approach is often expensive and energy-intensive. Monitoring oxygen levels under ice is a crucial tool for assessing the health of fish populations and informing management decisions.

In conclusion, oxygen depletion is a central and often lethal consequence of lake freezing. Understanding the factors that contribute to oxygen depletion, the varying oxygen requirements of different fish species, and the effectiveness of mitigation strategies is essential for managing freshwater ecosystems and preserving fish populations in cold climates. The interplay between ice cover, organic matter decomposition, and photosynthetic activity creates a complex dynamic that necessitates careful monitoring and informed management practices to minimize the risk of winterkill and ensure the long-term health of these aquatic environments.

3. Temperature Stratification

During the freezing process, and subsequent ice cover, lakes often exhibit temperature stratification. Water reaches its maximum density at approximately 4C. Consequently, as surface waters cool towards freezing, the colder, less dense water remains at the surface, eventually forming ice. The slightly warmer, denser water (around 4C) sinks to the bottom, creating a temperature gradient within the water column. This stratification is critical to fish survival. The deeper, warmer water provides a refuge from the frigid surface temperatures and prevents the entire lake from freezing solid. This denser bottom layer of water can maintain a relatively stable temperature, offering a more habitable environment for fish that are adapted to colder conditions but still require temperatures above freezing to survive. For example, during extended periods of ice cover, fish species like lake trout congregate in these deeper, slightly warmer zones.

The degree of temperature stratification is influenced by factors such as lake depth, morphology, and the duration of ice cover. Deeper lakes tend to exhibit more pronounced stratification than shallow lakes. Furthermore, the presence of springs or groundwater inflows can disrupt the stratification and influence water temperature distribution. The stratification also affects nutrient cycling and oxygen distribution within the lake. The warmer bottom layer can support microbial activity that consumes oxygen, potentially exacerbating oxygen depletion in that zone. Conversely, the ice cover prevents wind-driven mixing, limiting oxygen replenishment from the atmosphere. Understanding the dynamics of temperature stratification is thus vital for predicting oxygen levels and assessing the potential for winterkill events.

In summary, temperature stratification is a fundamental aspect of how lakes respond to freezing conditions and directly affects the survival of fish. The formation of a warmer, denser bottom layer provides a crucial thermal refuge. However, this stratification also creates complex interactions related to oxygen availability and nutrient cycling, influencing the overall health and stability of the aquatic ecosystem. Effective management strategies must consider temperature stratification and its implications for fish habitat and water quality under ice.

4. Antifreeze Proteins

The survival of numerous fish species in sub-zero aquatic environments depends critically on the presence and function of antifreeze proteins (AFPs). These specialized proteins prevent the formation and propagation of ice crystals within the body fluids of fish, thereby mitigating cellular damage and ensuring physiological function at temperatures below the freezing point of water. The production of AFPs is a primary adaptation enabling fish to inhabit lakes that freeze during winter.

  • Mechanism of Action

    AFPs function by binding to the surface of ice crystals, inhibiting their growth and preventing them from aggregating into larger, more damaging structures. They do not prevent freezing outright but rather control the ice crystal formation process. The binding is highly specific and varies depending on the structure of the AFP. Certain AFPs may bind more strongly to specific crystallographic planes of ice, further influencing the shape and size of the crystals formed.

  • Diversity of Types

    AFPs exhibit considerable structural diversity. Several distinct classes of AFPs have been identified, including alanine-rich alpha-helical proteins, cysteine-rich globular proteins, and carbohydrate-containing AFGPs (antifreeze glycoproteins). The specific type of AFP produced varies among fish species and may be related to the severity of the freezing conditions they typically encounter. For example, Antarctic fish possess particularly potent AFPs to survive in the extremely cold Southern Ocean.

  • Seasonal Production

    AFP production is often seasonally regulated, increasing during the fall and winter months in response to declining water temperatures. This upregulation is triggered by environmental cues such as decreasing photoperiod and temperature, which activate gene expression pathways responsible for AFP synthesis. As water temperatures rise in the spring, AFP production typically decreases, reflecting a reduced need for cryoprotection.

  • Ecological Significance

    The presence of AFPs allows fish to maintain activity and feeding behavior during winter, granting them a competitive advantage over species lacking such adaptations. The distribution and abundance of AFP-producing fish are often correlated with the severity and duration of ice cover in their respective habitats. The ability to resist freezing damage is crucial for survival in these environments and shapes community structure and ecological interactions.

The adaptive significance of AFPs is underscored by the prevalence of these proteins in fish inhabiting lakes subject to freezing. Their existence and function represent a critical evolutionary response to the challenges posed by sub-zero environments, allowing fish to thrive despite the inherent risks associated with ice formation. Without AFPs, many fish species would be unable to survive the winter months in temperate and arctic lakes. The study of AFPs continues to provide valuable insights into the physiological mechanisms underlying cold adaptation and the ecological dynamics of freshwater ecosystems.

5. Behavioral Adaptations

As lake ice forms, fish adjust their behavior to contend with decreased temperatures, limited oxygen, and reduced light. These behavioral shifts are essential for minimizing energy expenditure and maximizing survival prospects under challenging environmental conditions.

  • Reduced Activity and Torpor

    One of the primary behavioral responses is a significant reduction in activity levels. Fish enter a state of torpor, minimizing movement to conserve energy reserves. This quiescence reduces metabolic demand, allowing individuals to survive on limited oxygen supplies. For example, some species congregate in deeper, colder zones where activity is naturally suppressed. This behavioral modification can be seen as an adaptation to scarce resources and increased physiological stress.

  • Habitat Selection and Aggregation

    During winter, fish often congregate in specific areas of the lake that offer favorable conditions. These areas may include deeper zones with slightly warmer temperatures or locations near springs or groundwater inflows that provide higher oxygen levels. Such aggregation behavior increases the likelihood of finding suitable microhabitats and potentially reduces predation risk. For instance, certain fish species will gather around submerged structures or vegetation, utilizing these as refugia.

  • Dietary Changes

    The availability of food resources changes significantly under ice. Many fish species reduce or cease feeding entirely, relying on stored energy reserves. Others may switch to alternative food sources that are more readily available during winter, such as detritus or small invertebrates. These dietary shifts reflect an adaptation to seasonal resource scarcity and are crucial for maintaining energy balance during the lean winter months. This adaptability is seen in species that shift from insectivory during summer to consuming available plant matter in winter.

  • Vertical Migration

    Some fish species undertake vertical migrations within the water column to locate more favorable temperature or oxygen conditions. These movements can be influenced by factors such as temperature stratification or localized oxygen depletion. Fish may move to deeper zones during the day to avoid predation and then ascend to shallower areas at night to feed, if conditions allow. This type of migration demonstrates an active response to the dynamic environmental conditions under ice.

Collectively, these behavioral adaptations are integral to the overwintering success of fish populations in lakes subject to freezing. By reducing activity, selecting suitable habitats, modifying diets, and undertaking migrations, fish enhance their ability to survive the harsh conditions imposed by ice cover. The specific suite of behaviors exhibited varies among species and is influenced by the unique characteristics of each lake environment. Understanding these behavioral responses is crucial for predicting the impacts of climate change and managing freshwater fisheries in cold regions.

6. Ice Cover Duration

The length of time a lake remains frozen profoundly influences the aquatic ecosystem and the survival of fish populations. The duration of ice cover directly affects light penetration, water temperature, and oxygen availability, thereby modulating physiological and behavioral responses in fish. Extended ice cover can exacerbate existing stressors, while shorter periods may lessen the overall impact on the fish community.

  • Oxygen Depletion Amplification

    Longer ice cover periods prevent atmospheric oxygen exchange, leading to prolonged oxygen depletion. Microbial decomposition of organic matter continues to consume oxygen, and if the ice persists for an extended duration, oxygen levels can reach critically low levels, resulting in widespread fish mortality, known as winterkill. Lakes with lengthy ice cover are, therefore, at greater risk of experiencing these oxygen-related mortality events.

  • Light Limitation and Primary Production

    Extended ice cover limits light penetration, inhibiting photosynthesis by aquatic plants and algae. Reduced primary production diminishes the oxygen supply in the water and limits food availability for herbivorous fish and invertebrates. The longer the ice cover, the more significant the reduction in primary productivity, impacting the entire food web. For example, with reduced light penetration, the growth rates of juvenile fish dependent on visual foraging can be severely impacted.

  • Temperature Stability and Habitat Compression

    The persistence of ice cover helps maintain relatively stable, though cold, water temperatures. However, this can lead to habitat compression as fish seek refuge in limited areas with slightly warmer temperatures or higher oxygen concentrations. Overcrowding in these refuge areas can increase competition for resources and elevate the risk of disease transmission. The longer the ice cover, the greater the potential for habitat compression and associated stressors.

  • Reproductive Cycle Disruption

    Ice cover duration can disrupt the reproductive cycles of fish species that spawn in early spring. Prolonged ice cover may delay spawning or reduce the availability of suitable spawning habitat. This disruption can lead to reduced recruitment and impact the long-term population dynamics of affected species. The timing of ice melt is critical for many spring-spawning fish, and any significant shift in ice cover duration can have cascading effects on their reproductive success.

In conclusion, the amount of time that ice covers a lake is a significant determinant of the conditions faced by its fish populations. The length of this period influences oxygen levels, light availability, temperature stability, and reproductive success, impacting the overall health and survival of fish. An understanding of the connection between ice cover duration and fish survival is critical for effective fisheries management and conservation efforts in regions subject to seasonal freezing, particularly given the impacts of climate change on ice dynamics.

7. Habitat Selection

During periods of lake freezing, habitat selection becomes a paramount determinant of fish survival. The physical and chemical alterations induced by ice cover impose significant constraints, compelling fish to actively seek out areas that offer the most favorable conditions for overwintering.

  • Thermal Refugia Selection

    Fish often congregate in deeper regions or near groundwater inflows, which maintain slightly warmer temperatures compared to surface waters. These thermal refugia provide a critical buffer against the frigid conditions, reducing metabolic stress and energy expenditure. For example, lake trout and burbot are known to seek out such areas, where the water temperature remains above freezing, allowing them to conserve energy and maintain essential physiological functions. The availability and accessibility of these refugia directly influence overwinter survival rates.

  • Oxygenated Zone Preference

    Oxygen depletion is a common consequence of ice cover. Consequently, fish actively seek out areas with higher dissolved oxygen concentrations. These may include locations near springs, where oxygen-rich groundwater enters the lake, or areas with greater water flow. Species like trout, which have high oxygen requirements, are particularly reliant on finding these oxygenated zones. Failure to locate adequate oxygenated habitat can lead to physiological stress and, ultimately, mortality. The size and accessibility of these oxygenated zones become limiting factors for overwinter survival.

  • Shelter and Predation Avoidance

    Under-ice environments can alter predator-prey dynamics. Fish often seek shelter in submerged vegetation, rocky structures, or benthic zones to reduce predation risk. Reduced visibility due to ice and snow cover can make fish more vulnerable, increasing the importance of selecting habitats that offer protection. For instance, small forage fish may aggregate within dense macrophyte beds to avoid predation by larger piscivorous species. Habitat selection thus reflects a trade-off between finding favorable environmental conditions and minimizing the risk of being preyed upon.

  • Reduced Light Habitats

    Ice and snow cover significantly reduce light penetration into the water column, impacting fish behavior and distribution. While some species may prefer darker conditions to reduce predation risk, others require sufficient light for foraging. Fish species like walleye, which are adapted to low-light environments, might exhibit a competitive advantage under ice cover. The selection of habitats with appropriate light levels is therefore critical for maintaining foraging efficiency and energy balance during the winter months.

The act of choosing a habitat is influenced by both the physiological needs of the species and the environmental conditions under the ice. The success of a fish in overwintering often depends on its capacity to identify and locate areas that offer an optimal balance of temperature, oxygen, shelter, and light. The relationship between survival and habitat selection becomes especially important as lakes respond to the changing climate.

8. Winterkill Risk

When surface waters freeze, the risk of winterkill escalates, representing a significant threat to fish populations in temperate and arctic lakes. This phenomenon is a direct consequence of the environmental alterations imposed by ice cover, impacting oxygen levels, light penetration, and water temperature, all of which critically influence fish survival. The interplay of these factors determines the severity and extent of winterkill events.

  • Oxygen Depletion Under Ice

    Ice cover prevents atmospheric oxygen from dissolving into the water, while decomposition processes continue to consume dissolved oxygen. The longer the ice persists, the more oxygen is depleted. Fish require oxygen for respiration, and as levels drop below critical thresholds, they experience stress and eventually suffocate. Shallow lakes with high organic matter content are particularly susceptible to severe oxygen depletion, increasing the likelihood of winterkill. For instance, a shallow eutrophic lake covered in thick ice and snow for several months will likely experience near-anoxic conditions, resulting in substantial fish mortality.

  • Snow Cover Exacerbation

    Snow accumulation on the ice further reduces light penetration, inhibiting photosynthesis by aquatic plants and algae. This reduction in photosynthetic activity diminishes oxygen production, exacerbating oxygen depletion. Snow cover intensifies the effects of ice cover, accelerating the decline in oxygen levels and shortening the timeframe for winterkill to occur. For example, a lake with heavy snowfall during winter will experience a more rapid decline in oxygen than a lake with clear ice and minimal snow.

  • Species-Specific Vulnerability

    Different fish species exhibit varying tolerances to low oxygen conditions. Species with high oxygen demands, such as trout and salmon, are more vulnerable to winterkill than species with lower oxygen requirements, such as carp and bullheads. The composition of the fish community in a lake influences its susceptibility to winterkill events. For instance, a lake dominated by trout populations will experience more pronounced impacts from winterkill compared to a lake with a diverse assemblage of fish species with varying oxygen tolerances.

  • Delayed Mortality and Population Impacts

    Winterkill does not always result in immediate and complete fish mortality. Prolonged exposure to low oxygen conditions can weaken fish, making them more susceptible to disease and predation. This delayed mortality can further reduce fish populations in the months following ice melt. Furthermore, winterkill events can alter the age structure and genetic diversity of fish populations, potentially impacting their long-term resilience. For instance, a severe winterkill event may disproportionately affect older, larger fish, leading to a decline in reproductive output and genetic diversity.

Winterkill represents a significant ecological challenge in regions with seasonal freezing. The factors contributing to winterkill, including oxygen depletion, snow cover, species vulnerability, and delayed mortality, highlight the complex interactions within frozen aquatic ecosystems. Understanding these interactions is vital for managing freshwater fisheries and mitigating the impacts of winterkill events, particularly in the context of climate change and altered ice dynamics.

9. Light Reduction

The formation of ice on a lake surface precipitates a significant reduction in light penetration into the water column. This light reduction is a direct consequence of the physical properties of ice and snow, which scatter and absorb incoming solar radiation. Clear ice itself diminishes light transmission, but the presence of snow cover on the ice surface dramatically exacerbates this effect. The decreased light availability impacts primary productivity, alters fish behavior, and influences predator-prey interactions. The severity of light reduction depends on ice thickness, snow depth, and the presence of impurities within the ice. For example, a thick layer of snow-covered ice can effectively block almost all sunlight from reaching the water below, creating near-total darkness.

The diminished light availability significantly curtails photosynthetic activity by aquatic plants and algae. This reduction in primary production diminishes the oxygen supply in the water, further compounding the problem of oxygen depletion caused by ice cover. The decreased light can also affect the visual acuity of fish, impacting their ability to forage effectively and avoid predators. Some fish species adapt by shifting their activity patterns or altering their diet to accommodate the reduced light conditions. For instance, nocturnal species may become more active during the day, and fish that rely on visual hunting may switch to alternative feeding strategies, such as consuming detritus. Juvenile fish are especially vulnerable to light reduction, as their growth and survival depend on sufficient food resources and effective predator avoidance.

In summary, light reduction is a key element of the suite of environmental changes that accompany lake freezing. It initiates a cascade of ecological effects, impacting primary productivity, oxygen levels, and fish behavior. An understanding of light reduction is crucial for predicting the impacts of ice cover on freshwater ecosystems and for developing effective management strategies to mitigate the effects of winterkill and support fish populations during the winter months. The long-term effects of climate change on ice dynamics and light availability are a growing concern for fisheries managers and conservationists, necessitating further research and monitoring efforts.

Frequently Asked Questions

This section addresses common questions regarding the effects of lake freezing on fish populations, providing concise and informative answers based on current scientific understanding.

Question 1: How do fish survive when a lake freezes over?

Fish employ several strategies to survive under ice, including reducing their metabolic rate to conserve energy, seeking out warmer or more oxygenated areas, and producing antifreeze proteins to prevent ice crystal formation in their tissues.

Question 2: What is winterkill, and what causes it?

Winterkill is a phenomenon where fish die due to oxygen depletion under ice cover. Decomposition of organic matter consumes oxygen, while ice prevents atmospheric replenishment. Reduced light from snow cover further inhibits oxygen production by aquatic plants.

Question 3: Do all fish species survive equally well under ice?

No. Different species have varying tolerances to cold temperatures and low oxygen levels. Species like trout and salmon are more susceptible to winterkill than species like carp and bullheads, which are more tolerant of low oxygen conditions.

Question 4: How does temperature stratification affect fish during winter?

Temperature stratification creates distinct layers of water with varying temperatures. Fish often seek refuge in the slightly warmer, denser water near the bottom of the lake, which provides a thermal buffer against the frigid surface temperatures.

Question 5: Can human activities influence fish survival in frozen lakes?

Yes. Nutrient pollution from agricultural runoff or sewage can increase organic matter decomposition, exacerbating oxygen depletion and increasing the risk of winterkill. Proper land management practices can help reduce nutrient loading and protect fish populations.

Question 6: How does climate change impact fish survival in frozen lakes?

Climate change can alter ice cover duration and snow accumulation patterns, affecting water temperature, light penetration, and oxygen availability. These changes can disrupt fish reproductive cycles, increase the frequency and severity of winterkill events, and alter the distribution and abundance of fish species.

Understanding the responses of fish to lake freezing and the factors that influence their survival is crucial for fisheries management and conservation efforts.

Further research and monitoring are essential to assess the long-term impacts of environmental changes on these delicate aquatic ecosystems.

Survival Strategies for Fish Under Ice

Understanding the challenges faced by fish when a lake freezes is vital for effective conservation and management. These tips provide insights into factors influencing fish survival in frozen environments.

Tip 1: Understand Oxygen Dynamics: Recognize that ice and snow cover impede atmospheric oxygen exchange. Monitor oxygen levels, particularly in shallow, eutrophic lakes, as these are prone to depletion.

Tip 2: Protect Littoral Zones: Maintain healthy littoral zones as they provide refuge and feeding grounds. Submerged vegetation offers shelter and localized oxygen production.

Tip 3: Minimize Nutrient Loading: Reduce nutrient runoff from agricultural and urban sources. Excess nutrients fuel algal blooms, which consume oxygen during decomposition, exacerbating winterkill risks.

Tip 4: Consider Species-Specific Vulnerabilities: Acknowledge that certain fish species (e.g., trout, salmon) are more sensitive to low oxygen levels than others (e.g., carp, bullheads). Tailor management strategies to protect vulnerable species.

Tip 5: Monitor Ice Cover Duration: Track the duration of ice cover, as prolonged periods can intensify oxygen depletion and limit light penetration, impacting primary productivity.

Tip 6: Maintain Habitat Diversity: Ensure a variety of habitat types within the lake ecosystem. Deep zones, springs, and vegetated areas can provide thermal refugia and oxygenated zones.

Tip 7: Address Climate Change Impacts: Recognize that climate change is altering ice dynamics. Develop management plans that account for changing ice cover patterns and their effects on fish populations.

Implementing these strategies will help mitigate the negative consequences of ice cover and improve the long-term survival prospects for fish populations. Protecting these valuable resources requires a proactive and informed approach.

The subsequent conclusion will summarize the major findings and offer suggestions for future research and action.

Conclusion

This exploration of “what happens to fish when a lake freezes” reveals a complex interplay of environmental factors that directly impact survival. Ice cover induces significant alterations in oxygen availability, temperature stratification, and light penetration, compelling fish to employ various physiological and behavioral adaptations. Prolonged ice duration, coupled with snow cover, heightens the risk of winterkill, particularly in shallow, nutrient-rich lakes. Habitat selection becomes crucial as fish seek thermal refugia and oxygenated zones, but limited resources can lead to habitat compression and increased competition. The consequences for fish populations can be severe, potentially leading to population declines, altered community structure, and long-term ecological impacts.

The information presented underscores the importance of understanding the ecological consequences of lake freezing, particularly in the context of a changing climate. Further research should focus on species-specific responses to these challenging conditions and the efficacy of various mitigation strategies. Effective management practices must consider the interplay of factors influencing fish survival under ice to protect freshwater ecosystems and ensure the long-term health and resilience of fish populations in these fragile environments. Action is needed to address nutrient pollution and minimize anthropogenic stressors impacting these valuable aquatic resources.