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Supercooling Capacity and Cryoprotectants of Overwintering Larvae from Different Populations of Holcocerus Hippophaecolus

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Supercooling Capacity and Cryoprotectants of Overwintering Larvae from Different Populations of Holcocerus Hippophaecolus CryoLetters 37 (3), 206-217 (2016) © CryoLetters, [email protected] SUPERCOOLING CAPACITY AND CRYOPROTECTANTS OF OVERWINTERING LARVAE FROM DIFFERENT POPULATIONS OF HOLCOCERUS HIPPOPHAECOLUS Bin Tian, Lili Xu, Miao Zhang, Yuqian Feng and Shixiang Zong* Beijing Key Laboratory for Forest Pest Control, Beijing Forestry University, Beijing, 100083, P.R. China *Corresponding author e-mail: [email protected] Abstract BACKGROUND: Holcocerus hippophaecolus is the most serious pest occurred in seabuckthorn forest of three north areas. OBJECTIVE: The primary aims of the current study were to explore the physiological mechanisms and adaptability of H. hippophaecolus to low temperatures. MATERIALS AND METHODS: Assessing supercooling point, freezing point, and cryoprotectants of different larval instars from three different populations. RESULTS: Supercooling capacity of larvae from the 8–13 instar groups was relatively independent of temperature and other indicators such as latitude. Larvae from the 14–16 instar groups were sensitive to temperature and latitude, with generally lower limits and a wider range of SCPs than those of the other instar groups. CONCLUSION: For each population, the differences in the supercooling capacity of different instar stages for the identical period were not significant. The metabolism of fat and glycogen might not be the primary factors affecting the supercooling capacity. Keywords: Holcocerus hippophaecolus, overwintering larvae, different populations, different instars, supercooling point, cryoprotectant content INTRODUCTION reported by Bachmetjew as discussed by Somme (4). The supercooling point (SCP) of an insect is Insects are poikilothermic animals, and an important indicator of cold hardiness and a climate change is a primary factor that can strategy for overwintering and has been widely influence seasonal growth and cause declines in used to predict potential dispersal and populations. In both temperate and frigid geographic distributions of insect source regions, insects experience annual cold stress. populations. Therefore, cold hardiness is an important factor The supercooling capacity of insects is in the life history of insects in these areas, highly variable between both individuals and affecting reproduction, dispersal, distribution, groups, and the primary influences on and population dynamics in the following supercooling capacity are divided into external seasons (1) To adapt to low environmental and internal factors. The external factors temperatures, insects developed strategies to primarily include seasonal change, geographic safely overwinter in the long-term process of condition, and host-plant species, among others, evolution. Maintaining a supercooled state is a whereas the internal factors include the primary strategy for insects to overwinter in the biological state, developmental stage, and temperate and frigid regions of the Northern physiological and metabolic state of the body. Hemisphere (2). Additionally, the supercooling point in insects is Fahrenheit (1842) discovered supercooling affected by a change in the water ratio with the and found that water remains a liquid when the production and transformation of temperature falls below the freezing point (FP) cryoprotectants, particularly fats, low-molecular under certain conditions reported by Li (3). The weight substances, and polyhydric alcohols, phenomenon of supercooling in insects was first amino acids, and anti-freezing proteins (AFPs), 206 which are stimulated by changes in environment, will provide the theoretical basis to assess host, developmental stage, and biological state. overwintering strategies, bio-physiological The interaction of these external and internal regulation, and the potential for population factors regulates the supercooling capacity of establishment. insects (2). In the long life history of Holcocerus MATERIALS AND METHODS hippophaecolus (Lepidoptera: Cossidae), 4 years are required to complete a generation, which Study sites and sample collection. commonly has 16 instars (5). The larva feeds on The specimens used in these experiments the trunk and roots of sea buckthorn. Newly were collected from the field in China during hatched larvae bore primarily into the phloem of mid-March 2013. The three different collection the trunk, which leads to desiccation of the bark. sites were Jianping in Liaoning Province (JP), A few larvae bore into the xylem and damage Zhungeerqin in Inner Mongolia (ZQ), and roots as they move underground to overwinter, Pengyang in Ningxia Hui Autonomous Region and, as the damage accumulates and most roots (PY; Table 1). All overwintering larvae were are hollowed out, the host eventually dies (6-10). transported in soil with roots of sea buckthorn to In recent years, most areas with sea buckthorn the laboratory at Beijing Forestry University experienced outbreaks of this species, including within 24 h without any treatment; the healthy the provinces of Shaanxi, Shanxi, Liaoning, and larvae were transferred to the laboratory for Hebei and Inner Mongolia and Ningxia Hui testing. The local temperature (°C) data were autonomous regions. The damaged area has obtained from the China Meteorological Data increased to 133,000 hm2, with over 67,000 hm2 Sharing Service System (http:// cdc.cma.gov.cn). in which the mortality of sea buckthorn is total. Table 1. Geographic location, altitude, and temperature conditions at the collection sites. Collection Longitude Latitude Altitude Mean Jan. Mean Mar. Accumulated Low Site (°E) (°N) (m) Temp. (°C) Temp. (°C) Temp. (Days -°C) JP 119°54' 41°19' 441 -10.6 1.5 -952.3 ZQ 110°25' 39°52' 1169 -7.4 4.4 -798.0 PY 106°16' 35°50' 1788 -5.3 7.3 -455.2 The accumulated low temperature is the mean of the total subzero daily temperatures from November 2012 to March 2013. JP, JianPing in Liaoning Province; ZQ, Zhungeerqi in Inner Mongolia Autonomous Region; PY, Pengyang in Ningxia Hui Autonomous Region. Moreover, damage caused by this species is Measurement of instars. increasing. The general research on H. All larvae were photographed under a hippophaecolus has concentrated primarily on microscope (LEICA EZ 4D). The head widths bio-ecological characteristics, population and body lengths of instars were measured, with dynamics, chemical prevention, sex the magnification noted(6). Based on the pheromones, mechanisms to explain outbreaks, distribution of measurements, three groups of and monitoring for early warning of outbreaks instars (8–10, 11–13, and 14–16) were tested. (11-18). Although the collective achievements are considerable, the cold hardiness of H. Measurement of SCP and FP. hippophaecolus from different populations has Ten or more larvae of each group of instars not been investigated. from each population were selected. Each larva The primary aims of the current study were was fixed to a thermocouple probe with to explore the physiological mechanisms and parafilm, and the thermocouple was connected adaptability of H. hippophaecolus to low to an automatic data recorder (an instrument to temperatures by assessing the supercooling measure the SCP). A larva was placed in a capability of different groups of larval instars cotton-lined container and transferred into a from three populations. The results of this study freezing chamber (high-low temperature test 207 chamber), and, at the cooling rate of 1°C/min, Measurement of glycogen content. the body temperature of the larva was recorded. Glycogen content was determined using the The temperature at which an exothermic phenol and concentrated sulfuric acid method response was first detected was the supercooling (20). Each larva was dried, homogenized in 10 point, and the FP was the point at which larval ml tubes with 70% ethanol, and then centrifuged temperature continued to decline following at 2,600 × g for 10 min three times before conclusion of the exothermic process. recovering the supernatant (21). The residue was mixed with 6 ml of 10% (vol:vol) trichloroacetic Measurement of water content. acid and heated at 70°C for 15 min in an To assess the water content, ten more larvae electrothermostatic water bath. The mixture were selected from each of the three groups of cooled and was then centrifuged for 15 min at instars from each of the three populations. The 3,000 × g. A UV spectrophotometer was used to fresh mass (FM) of an individual larva was determine the absorbance values of supernatant determined on an electronic balance (±0.001 g). extracts at 490 nm. Measurements were obtained The larva was then oven-dried at 60°C for 72 h using 5–10 larvae from each population of H. to determine dry mass (DM). The water content hippophaecolus. as a ratio was calculated using FW and DW in the following formula: (FM − DM)/FM × 100%. Statistical analyses. All statistical analyses were conducted Measurement of fat content. using the SPSS 19.0.0 statistical software The fat content was measured as described package. The mean SCPs and FPs and water, fat, by Liu et al. A dried larva was homogenized, and glycogen contents were analyzed and and the fat content was extracted using a mixture compared with one-way ANOVA (α = 0.05) and of chloroform and methanol (19) in the least significant difference tests (LSD, α = 0.05). following steps. A dried larva was ground to Pearson’s correlations were used to assess the powder in a mortar. The powdered, dried larva relationships between the SCPs and FPs and the was weighed (DW), placed in a 10 ml tube, and geographical, climatic, and physical data. homogenized
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