Interactive effects of soil warming and fertilization on root production, mortality, and longevity in a Norway spruce stand in Northern Sweden

The effects of soil warming and nitrogen availability on root production, longevity and mortality were studied using minirhizotrons in irrigation (C), fertilized (F), heated (H), and heated‐fertilized (HF) plots in a Norway spruce stand in northern Sweden from October 1996 to October 1997. Irrigation was included in all treatment plots. Heating cables were used to maintain the soil temperature in heated plots at 5°C above that in unheated plots during the growing season. A Kaplan–Meier approach was used to estimate the longevity of fine roots and Cox proportional hazards regression to analyze the effects of the H, F, and HF treatments on the risk of root mortality. The proportion of annual root length production contributed by winter–spring production amounted to 52% and 49% in heated plots and heated‐fertilized plots, respectively. The annual root length production in C plots was significantly higher than in other treatments, while the HF treatment gave significantly greater production compared with the F treatment. The risk of mortality (hazard ratio) relative to C plots was higher in H plots (358%) and F plots (191%). The interaction between heating and fertilizing was strongly significant. The increase in the risk of root mortality in combined fertilization and heating (103%) was lower than that in the H or F plots. The results show that nitrogen addition combined with warmer temperatures decreases the risk of root mortality, and fine root production is a function of the length of the growing season. In the future, fertilization combined with the warmer temperatures expected to follow predicted climatic change may increase root production in boreal forests at low fertility sites.     

Experimental soil warming effects on CO2 and CH4 flux from a low elevation spruce–fir forest soil in Maine, USA

The effect of soil warming on CO₂ and CH₄ flux from a spruce–fir forest soil was evaluated at the Howland Integrated Forest Study site in Maine, USA from 1993 to 1995. Elevated soil temperatures (～5 °C) were maintained during the snow-free season (May – November) in replicated 15 × 15-m plots using electric cables buried 1–2 cm below the soil surface; replicated unheated plots served as the control. CO₂ evolution from the soil surface and soil air CO₂ concentrations both showed clear seasonal trends and significant (P < 0.0001) positive exponential relationships with soil temperature. Soil warming caused a 25–40% increase in CO₂ flux from the heated plots compared to the controls. No significant differences were observed between heated and control plot soil air CO₂ concentrations which we attribute to rapid equilibration with the atmosphere in the O horizon and minimal treatment effects in the B horizon. Methane fluxes were highly variable and showed no consistent trends with treatment.     

Effects of soil warming during spring on photosynthetic recovery in boreal Norway spruce stands

The effect of soil thawing and soil temperature on postwinter recovery of photosynthetic capacity was studied, during late spring and early summer, in Norway spruce stands in northern Sweden. Soil temperature was manipulated by means of buried heating cables. The warming treatment was applied to stands with low (natural) and high (fertilized) availability of nutrients. Soil thawing, expressed as water availability, was followed by means of sapflow in stems, and shoot water potentials. The recovery of photosynthetic capacity was assessed by measuring the rate of light-saturated photosynthesis (A_max), and maximum photochemical efficiency of photosystem II in detached shoots, and chlorophyll a fluorescence. Accumulation of starch reserves in the needles was followed as an independent indicator of photosynthetic performance in situ. Snowmelt and soil thawing occurred more than one month earlier in heated than in unheated plots. This was expressed both as sapflow and as differences in shoot water potential between treatments. During May, the rates of A_max were significantly higher on heated than on control plots. The effect of soil warming on A_max was, however, not reflected in chlorophyll fluorescence or needle starch content. The time course of the recovery of photosynthetic capacity was mainly controlled by mean air temperature and by the frequency of severe night frosts, and to a lesser extent by earlier soil thawing and higher soil temperatures.     

The effect of root temperature on rose plants in relation to air temperature

Rose plants (Rosa hybrida ‘Sonia’=‘Sweet Promise’) were grown in heated (minimum night temperature 17°C), and unheated greenhouses with or without root heating to 21°C. These trials covered 6 growth cycles extending over two winter seasons. In the heated greenhouse, root heating did not increase yield, flower quality or plant development. In the unheated greenhouse, root-heated plants grew as well as those in the air-heated greenhouse as long as the air temperature did not fall below 6°C. When minimum night temperatures fell below 6°C, growth, yield and quality were reduced, irrespective of root temperature. Daytime plant water relations were studied in plants growing at 6 different root temperatures in the unheated greenhouse. Leaf resistance to water diffusion was lowest at optimal root temperature. Total leaf water potential was not significantly affected by root temperature.     

Infrared heater system for warming tropical forest understory plants and soils

The response of tropical forests to global warming is one of the largest uncertainties in predicting the future carbon balance of Earth. To determine the likely effects of elevated temperatures on tropical forest understory plants and soils, as well as other ecosystems, an infrared (IR) heater system was developed to provide in situ warming for the Tropical Responses to Altered Climate Experiment (TRACE) in the Luquillo Experimental Forest in Puerto Rico. Three replicate heated 4‐m‐diameter plots were warmed to maintain a 4°C increase in understory vegetation compared to three unheated control plots, as sensed by IR thermometers. The equipment was larger than any used previously and was subjected to challenges different from those of many temperate ecosystem warming systems, including frequent power surges and outages, high humidity, heavy rains, hurricanes, saturated clayey soils, and steep slopes. The system was able to maintain the target 4.0°C increase in hourly average vegetation temperatures to within ± 0.1°C. The vegetation was heterogeneous and on a 21° slope, which decreased uniformity of the warming treatment on the plots; yet, the green leaves were fairly uniformly warmed, and there was little difference among 0–10 cm depth soil temperatures at the plot centers, edges, and midway between. Soil temperatures at the 40–50 cm depth increased about 3°C compared to the controls after a month of warming. As expected, the soil in the heated plots dried faster than that of the control plots, but the average soil moisture remained adequate for the plants. The TRACE heating system produced an adequately uniform warming precisely controlled down to at least 50‐cm soil depth, thereby creating a treatment that allows for assessing mechanistic responses of tropical plants and soil to warming, with applicability to other ecosystems. No physical obstacles to scaling the approach to taller vegetation (i.e., trees) and larger plots were observed.     

播种季节和种植方式对多年生苦荞主要农艺性状的影响

该研究选取六个多年生苦荞新品系,对春季、秋季直播与秋季再生其主要农艺性状进行调查。结果表明:(1)不同播种季节对多年生苦荞新品系主花序的花粉可育率、总结实率、有效结实率、植株株高、主茎粗、主茎分枝数、主茎节数、籽粒百粒重、单株粒数、单株产量的影响均达到显著或极显著水平;秋播主花序花粉可育率、总结实率、有效结实率、植株主茎分枝数、籽粒百粒重、单株粒数、单株产量均极显著高于春播;植株株高、主茎粗、主茎节数均极显著低于春播;主花序花朵大小、籽粒种子长宽比无显著差异。(2)不同种植方式对主花序花粉可育率、有效结实率、植株主茎节数及籽粒百粒重的影响达到显著或极显著水平;秋季再生主花序花粉可育率、籽粒单株粒数显著高于秋季直播;主花序有效结实率、植株主茎粗、主茎节数、籽粒百粒重显著低于秋季直播;主花序花朵大小、总结实率、植株株高、主茎分枝数、籽粒种子长宽比、单株产量无显著差异;相关分析表明,各生长季节下主花序有效结实率及单株粒数与单株产量的相关系数均最高。(3)所有参试品系中,1612-241秋季直播的单株产量显著高于其他品系; 1612-16、1612-33秋季再生单株产量较正季优势显著。该研究结果有助于筛选出适宜一季播种两季收获的优良品系,为今后多年生苦荞的选择育种提供线索基础。     

Changes in heated and autoclaved forest soils of S.E. Australia. II. Phosphorus and phosphatase activity

The effect of soil heat and autoclaving on labile inorganic P (Bray I), microbial P (P-flush) and on phosphatase activity was studied by heating five forest soils in the laboratory, which simulated the effects of heat during bushfires. Top soil was heated to 60 °C, 120 °C and 250 °C or autoclaved for 30 minutes. Soils were analysed immediately after heating and during seven months of incubation to assess immediate and longer-term effects of heating.Labile inorganic P increased immediately after heating and autoclaving soils, with the highest amount recorded for the 250 °C treatment. Phosphorus associated with microbial biomass decreased with heat, and none or small amounts were detected in soils heated to 250 °C and autoclaved, because high temperatures killed the microbial population. Most of the P released from microbes acted as a source of labile inorganic P in soils low in inorganic P, and some of the released P was fixed by the soil. In one soil high in inorganic labile P and with undetectable amounts of microbial-P, the increase in Bray P on heating could only be assigned to solubilisation of other sources of total P Because high temperatures denature enzymatic proteins, phosphatase activity diminished with the increase in temperature, and no activity was detected in 250 °C and autoclaved soils.Phosphorus released by heating decreased during incubation in three of the five soils studied, approaching values observed in unheated soils. Simultaneously, an increase in microbial P was observed in these heated soils, indicating that the partial recovery of microbial biomass acted as a sink for the decrease in Bray-P measured. Phosphatase activity recovered only partially during incubation of heated soils.     

Performance of High Arctic tundra plants improved during but deteriorated after exposure to a simulated extreme temperature event

Arctic ecosystems are known to be extremely vulnerable to climate change. As the Intergovernmental Panel on Climate Change scenarios project extreme climate events to increase in frequency and severity, we exposed High Arctic tundra plots during 8 days in summer to a temperature rise of approximately 9°C, induced by infrared irradiation, followed by a recovery period. Increased plant growth rates during the heat wave, increased green cover at the end of the heat wave and higher chlorophyll concentrations of all four predominating species (Salix arctica Pall., Arctagrostis latifolia Griseb., Carex bigelowii Torr. ex Schwein and Polygonum viviparum L.) after the recovery period, indicated stimulation of vegetative growth. Improved plant performance during the heat wave was confirmed at plant level by higher leaf photochemical efficiency (F_v/F_m) and at ecosystem level by increased gross canopy photosynthesis. However, in the aftermath of the temperature extreme, the heated plants were more stressed than the unheated plants, probably because they acclimated to warmer conditions and experienced the return to (low) ambient as stressful. We also calculated the impact of the heat wave on the carbon balance of this tundra ecosystem. Below‐ and aboveground respiration were stimulated by the instantaneous warmer soil and canopy, respectively, outweighing the increased gross photosynthesis. As a result, during the heat wave, the heated plots were a smaller sink compared with their unheated counterparts, whereas afterwards the balance was not affected. If other High Arctic tundra ecosystems react similarly, more frequent extreme temperature events in a future climate may shift this biome towards a source. It is uncertain, however, whether these short‐term effects will hold when C exchange rates acclimate to higher average temperatures.     

Response of the soil seed‐bank of Cumberland Plain Woodland to heating

Abstract Soil was investigated in a Cumberland Plain Woodland community to determine the presence of a soil seed‐bank and whether species richness and abundance of plants germinating from it were affected by heating such as that experienced in a fire. Soil samples were taken from the Holsworthy Military Area, in the south‐eastern region of the Sydney Basin, New South Wales, Australia, and one of four treatments was applied; soil was heated to 80°C, 40°C, unheated or unheated with litter not removed. Sixty‐eight species, representing 26 families including 11 exotic and 57 native species germinated from the soil. Herbs and grasses dominated and were in similar proportions to those surveyed in the above‐ground vegetation, suggesting that the soil seed‐bank reflected the current structure of the vegetation, although species composition differed. Species responded differently to heating. The seeds of some species germinated when heated at a higher temperature (80°C), particularly those from the family Fabaceae, whereas other species were more common in unheated or lightly heated samples (40°C). This suggests that fire is likely to change the species composition of the above‐ground vegetation and indicates that management must ensure that species that do not germinate when heated are maintained, as well as those species that germinate following heating. A large proportion of soil seed‐bank species showed low germination rates in the trials, and 112 above‐ground species did not germinate in the soil samples. We do not understand whether species of these two sets do not produce a soil‐stored seed‐bank or whether the seed‐bank has been depleted by past practices at Holsworthy. Further research is needed.     

Effects of warming on root diameter,distribution, and longevity in an alpine meadow

Roots form one of the most important carbon (C) pools in alpine ecosystems. Uncertainty about the effects of warming on root dynamics limits our ability to predict how C will transfer between biological and atmospheric pools in alpine regions under global warming. We used a minirhizotron technique to gain a better understanding of the response of alpine plant roots to warming. We looked for effects on root diameter, root depth in the soil, and root lifespan under a controlled asymmetrical warming (1.2/1.7 °C during daytime/nighttime) experiment during the growing season of 2009 in an alpine meadow on the northern Tibetan plateau. Roots became smaller in diameter, moved toward the upper soil layers, and showed significantly shorter lifespans in heated (H) than in unheated (UH) plots. Furthermore, in H treatment plots root lifespan was more strongly influenced by the time of root emergence rather than by root diameter. These results provide evidence that alpine plants may respond to climate change by altering their roots so that they are thinner, distributed shallower and turning over faster, suggesting that soil C flow and nutrient cycling would be accelerated resulting from the fast turnover of fine roots under elevated temperatures.     

Increased sensitivity of heat-stressed Saccharomyces cerevisiae cells to food-grade antioxidants

Unheated and heat-stressed Saccharomyces cerevisiae cells were examined for their relative sensitivities to butylated hydroxyanisole (BHA), tertiary butylhydroquinone (TBHQ), and propyl gallate. Heated cells had significant (P less than or equal to 0.05) increases in sensitivity to 50 micrograms of BHA, 100 micrograms of TBHQ, and 1,000 micrograms of propyl gallate per ml as compared with unheated cells when surface plated on antioxidant-supplemented recovery agar. The rate of increase in size of colonies developed by heated cells was slower than that of unheated cells, and the presence of antioxidants in recovery agar enhanced this effect. Heat-stressed cells also had increased sensitivity to ethanol. Incubation temperatures of 15, 21, 30, and 37 degrees C for enumerating unheated cells had no significant effect on the numbers of colonies formed on unsupplemented recovery agar; however, incorporation of 100 micrograms of BHA, 200 micrograms of TBHQ, or 1,000 micrograms of propyl gallate per ml into agar resulted in significant decreases in the number of colonies formed by heated cells at various incubation temperatures. The detrimental effects of TBHQ and propyl gallate on repair of heat-injured cells are apparently expressed at a temperature higher than that observed for BHA. It is suggested that the adverse effects of antioxidants on repair of heat-injured S. cerevisiae cells may be associated with oxygen availability.     

Increased sensitivity of heat-stressed Saccharomyces cerevisiae cells to food-grade antioxidants.

Unheated and heat-stressed Saccharomyces cerevisiae cells were examined for their relative sensitivities to butylated hydroxyanisole (BHA), tertiary butylhydroquinone (TBHQ), and propyl gallate. Heated cells had significant (P less than or equal to 0.05) increases in sensitivity to 50 micrograms of BHA, 100 micrograms of TBHQ, and 1,000 micrograms of propyl gallate per ml as compared with unheated cells when surface plated on antioxidant-supplemented recovery agar. The rate of increase in size of colonies developed by heated cells was slower than that of unheated cells, and the presence of antioxidants in recovery agar enhanced this effect. Heat-stressed cells also had increased sensitivity to ethanol. Incubation temperatures of 15, 21, 30, and 37 degrees C for enumerating unheated cells had no significant effect on the numbers of colonies formed on unsupplemented recovery agar; however, incorporation of 100 micrograms of BHA, 200 micrograms of TBHQ, or 1,000 micrograms of propyl gallate per ml into agar resulted in significant decreases in the number of colonies formed by heated cells at various incubation temperatures. The detrimental effects of TBHQ and propyl gallate on repair of heat-injured cells are apparently expressed at a temperature higher than that observed for BHA. It is suggested that the adverse effects of antioxidants on repair of heat-injured S. cerevisiae cells may be associated with oxygen availability.     

Effects of nutrition and soil warming on stemwood production in a boreal Norway spruce stand

The boreal forest is expected to experience the greatest warming of all forest biomes. The extent of the boreal forest, the large amount of carbon contained in the soil, and the expected climate warming, make the boreal forest a key biome to understand and represent correctly in global carbon models. It has been suggested that an increase in temperature could stimulate the release of CO₂ caused by an increased decomposition rate, more than biomass production, which could convert current carbon sinks into carbon sources. Most boreal forests are currently carbon sinks, but it is unclear for how long in the future the carbon sink capacity of the boreal forest is likely to be maintained. The impact of soil warming on stem volume growth was studied during 6 years, in irrigated (I) and irrigated‐fertilized (IL) stands of 40‐year‐old Norway spruce in Northern Sweden. From May to October heating cables were used to maintain the soil temperature on heated‐irrigated plots (Ih and ILh) 5 °C above that on unheated control plots (Ic and ILc). After six seasons' warming, stem volume production (m³ ha^?1 a^?1) was 115% higher on Ih than on unheated (Ic) plots, and on heated and irrigated‐fertilized plots (ILh) it was 57% higher than on unheated plots (ILc). The results indicate that in a future warmer climate, an increased availability of nitrogen, combined with a longer growing season, may increase biomass production substantially, on both low‐ and high‐fertility sites. It is, however, too early to decide whether the observed responses are transitory or long lasting. It is therefore crucial to gain a better understanding of the responses of boreal forest ecosystems to climate change, and to provide data to test and validate models used in predicting the impact of climate change.     

High night temperatures during grain number determination reduce wheat and barley grain yield: a field study

Warm nights are a widespread predicted feature of climate change. This study investigated the impact of high night temperatures during the critical period for grain yield determination in wheat and barley crops under field conditions, assessing the effects on development, growth and partitioning crop‐level processes driving grain number per unit area (GN). Experiments combined: (i) two contrasting radiation and temperature environments: late sowing in 2011 and early sowing in 2013, (ii) two well‐adapted crops with similar phenology: bread wheat and two‐row malting barley and (iii) two temperature regimes: ambient and high night temperatures. The night temperature increase (ca. 3.9 °C in both crops and growing seasons) was achieved using purpose‐built heating chambers placed on the crop at 19:000 hours and removed at 7:00 hours every day from the third detectable stem node to 10 days post‐flowering. Across growing seasons and crops, the average minimum temperature during the critical period ranged from 11.2 to 17.2 °C. Wheat and barley grain yield were similarly reduced under warm nights (ca. 7% °C^?1), due to GN reductions (ca. 6% °C^?1) linked to a lower number of spikes per m². An accelerated development under high night temperatures led to a shorter critical period duration, reducing solar radiation capture with negative consequences for biomass production, GN and therefore, grain yield. The information generated could be used as a starting point to design management and/or breeding strategies to improve crop adaptation facing climate change.     

Changes in heated and autoclaved forest soils of S.E. Australia. I. Carbon and nitrogen

The effect of heating and autoclaving on extractable nitrogen, N mineralisation and C metabolism was studied by heating five forest soils in the laboratory, simulating the range of effects of heat due to bushfire. Top soil (0–5 cm) was heated to 60 °C, 120 °C and 250 °C for 30 minutes; unheated soil was taken as a control. Samples of the soil heated to 250 °C were also inoculated with fresh soil to accelerate the recovery of the microbial population. Soil autoclaving was carried out as another heat treatment (moist heat). Soils were analysed immediately after heating and 3 times during seven months of incubation to assess immediate and longer-term effects of heating.Extractable N (organic and mineral forms) increased after heating to 120 °C, but decreased with further heating to 250 °C suggesting the volatilisation of N. N associated with microbial biomass diminished with heating and was barely detectable after the 250 °C treatment. Microbial biomass was an important source of soluble N in heated soils, and only partly recovered during subsequent long incubation. The amount of N mineralised during incubation depended on both soil and temperature. Nitrification did not occur when soils were heated to 250 °C (with or without inoculum), or after autoclaving, demonstrating the high sensitivity of nitrifiers to heat. At the beginning of soil incubation, respiration was enhanced in heated soils (250 °C, 250 °C inoculated) and autoclaved soils, but after 30 days of incubation respiration decreased to values either similar to or lower than those in control. This respiration pattern indicated that a fraction of labile C was released by heating, which was quickly mineralised within 30 days of incubation. These results demonstrate some effects of soil heating on C and N dynamics in forest soils.     

不同积温和种植密度对饲用黑麦分蘖动态的影响

应用回归均匀设计构造试验方案,采用分期播种的方法,研究了不同冬前积温、种植密度对饲用黑麦单株分蘖动态、群体分蘖动态以及有效茎数动态变化的影响,并建立了模拟模型。结果表明,饲用黑麦单株分蘖数量同时受播期、播量的影响,生长前期主要受播期早晚的影响,生长后期主要受播种密度的制约,总体来看,播期越早、播量越小,其单株分蘖数量就越多,反之就越少,饲用黑麦不论是在生长前期、中期,还是后期,随着播期推迟,各播量群体分蘖数不断下降;随着播量增大,各播期群体总蘖数不断上升,饲用黑麦群体有效茎数取决于播期和播量的共同作用,播期早,要控制播量不宜过大,合理的群体结构主要依靠单株有效茎数的增长潜力来保证;播期晚,要防止播量过小,合理的群体结构主要依靠主茎数量的增加来实现。     

Theory and performance of an infrared heater for ecosystem warming

In order to study the likely effects of global warming on future ecosystems, a method for applying a heating treatment to open-field plant canopies (i.e. a temperature free-air controlled enhancement (T-FACE) system) is needed which will warm vegetation as expected by the future climate. One method which shows promise is infrared heating, but a theory of operation is needed for predicting the performance of infrared heaters. Therefore, a theoretical equation was derived to predict the thermal radiation power required to warm a plant canopy per degree rise in temperature per unit of heated land area. Another equation was derived to predict the thermal radiation efficiency of an incoloy rod infrared heater as a function of wind speed. An actual infrared heater system was also assembled which utilized two infrared thermometers to measure the temperature of a heated plot and that of an adjacent reference plot and which used proportional–integrative–derivative control of the heater to maintain a constant temperature difference between the two plots. Provided that it was not operated too high above the canopy, the heater system was able to maintain a constant set-point difference very well. Furthermore, there was good agreement between the measured and theoretical unit thermal radiation power requirements when tested on a Sudan grass (Sorghum vulgare) canopy. One problem that has been identified for infrared heating of experimental plots is that the vapor pressure gradients (VPGs) from inside the leaves to the air outside would not be the same as would be expected if the warming were performed by heating the air everywhere (i.e. by global warming). Therefore, a theoretical equation was derived to compute how much water an infrared-warmed plant would lose in normal air compared with what it would have lost in air which had been warmed at constant relative humidity, as is predicted with global warming. On an hourly or daily basis, it proposed that this amount of water could be added back to plants using a drip irrigation system as a first-order correction to this VPG problem.     

极端晚播对小麦籽粒产量、氮素吸收利用和籽粒蛋白质含量的影响

为探索小麦高产高效优质生产技术途径,指导小麦晚播生产实践,2012年10月—2014年6月,以弱春性小麦偃展4110和半冬性小麦矮抗58为材料进行连续2年的田间定位试验,设置了常规适播(10月中旬、240万株·hm^-2)和极端晚播(11月中旬、600万株·hm^-2)两种栽培模式,研究了极端晚播对0～40 cm土层土壤硝态氮含量、小麦氮素吸收利用、产量、籽粒蛋白质含量和氮素吸收效率的影响.结果表明: 与常规适播处理相比,两个生长季极端晚播处理均使拔节和开花期0～40 cm土壤硝态氮含量显著提高,从而促进拔节后小麦植株氮素吸收积累,成熟期穗部氮素的分配比例也得到提高,最终显著提高小麦籽粒蛋白质含量和偃展4110的蛋白质产量、氮素吸收效率,但对籽粒产量的影响因品种而异.其中,极端晚播处理使偃展4110的籽粒产量显著提高,而矮抗58的籽粒产量却显著降低.因此,极端晚播栽培模式可维持小麦拔节后的土壤氮供应,有利于提高小麦氮素吸收效率,从而提高小麦籽粒产量和蛋白质含量,是灌区小麦高产优质的有效途径之一.     

The effect of sowing date and plant density on barley

Proctor and Maris Puma barleys were sown in October, early March, and late April at 50, 100, 200, 400 and 800 plants per m²and at low and high fertilizer levels. Shoot dry matter and grain dry matter showed no significant response to density for the first two sowings, but increased with increasing density in the last sowing. In all sowings the ear number per m²rose with increasing density, and grain number per ear fell, and there were only small interactions between density and sowing date, but the effect on weight per grain differed markedly in the last sowing from that in the first two sowings. Nitrogen concentration per cent of dry matter in the grain and in the straw showed little response to density and the values for both grain and straw were highest in the last sowing. Nitrogen content per m²for both shoot and grain at first rose, and then fell with increasing density. The maximum amount of nitrogen per m²was found at about 100 plants per m²in the early sowings, and at 400 plants per m²in the last sowing. The nitrogen data indicated a loss of nitrogen from the plant at high densities.