施氮对青藏高原东缘窄叶鲜卑花灌丛土壤呼吸的影响

开展土壤呼吸对大气氮沉降增加的响应研究对预测陆地生态系统碳循环具有重要意义。采用外施氮肥模拟氮沉降,结合壕沟法分离土壤呼吸组分,研究青藏高原东缘主要的灌丛类型——窄叶鲜卑花(Sibiraea angustata)灌丛土壤呼吸对不同施氮水平(N0(对照)、N2、N5和N10分别相当于0、2、5和10 g N m-2a-1浓度的氮沉降)的短期响应。结果表明:试验期间(2012年5—10月份),(1)土壤呼吸呈现明显的季节变化,施氮对生长季土壤总呼吸、异养呼吸无显著影响,而对自养呼吸有显著的抑制作用(P0.05)。(2)土壤呼吸也存在显著的日变化,施氮对一天中土壤总呼吸及其组分均有显著影响(P0.001)。总体上,施氮促进了土壤总呼吸、异养呼吸,而抑制了自养呼吸。(3)施氮对土壤总呼吸、异养呼吸平均每月排放CO2通量无显著影响,而对自养呼吸平均每月排放CO2通量有显著的抑制作用(P0.05),并在不同月份对土壤呼吸及其组分的影响不同。(4)土壤总呼吸、异养呼吸与地下5 cm土壤温度之间具有较好的指数关系(P0.001),而与土壤含水量相关性较弱。关于土壤呼吸各组分对大气氮沉降响应差异的机理有待进一步研究。     

模拟增温对高寒草甸植物叶片碳氮及其同位素δ~(13)C和δ~(15)N含量的影响

基于国际冻原计划(ITEX)模拟增温效应对植物影响的研究方法,以高寒矮嵩草草甸4种植物(矮嵩草、垂穗披碱草、棘豆、麻花艽)为实验材料,设置大(OTC1)和小(OTC2)两类增温小室,测定了其叶片碳氮及其稳定性碳同位素(δ~(13)C、δ~(15)N)等指标在增温3年后的变化。结果表明:除矮嵩草在较小增温小室(OTC2)中C/N比值比对照降低了14.1%,其它物种C/N在两个增温处理下都有所增加,但差异均不显著(P0.05)。4种植物叶片δ~(13)C值在-24.12‰~-28.34‰之间,矮嵩草叶片δ~(13)C值随增温而升高,棘豆、麻花艽随增温而降低,且矮嵩草在OTC2的δ~(13)C值变化达到显著水平(P0.05)。矮嵩草和麻花艽的叶片δ~(15)N值在OTC1和OTC2中均比对照增加,且麻花艽增加较显著(P0.05)。垂穗披碱草在OTC1和OTC2的叶片δ~(15)N值比对照分别减少18.7%和26.9%,差异都不显著(P0.05);棘豆叶片δ~(15)N值在OTC2内比对照低11.0%(P0.05),在OTC1的内比对照高2.8%(P0.05)。可见,高寒矮嵩草草甸不同功能群植物物种碳氮含量及稳定性碳氮同位素含量对短期增温有不同的响应模式和规律。     

胡杨(Populus euphratica)叶片呼吸作用对地下水埋深的响应

土壤水分条件能够改变叶片呼吸作用, 进而影响叶片、个体及生态系统的碳平衡。在内蒙古自治区额济纳旗境内, 根据地下水埋深度的差异建立了干旱和湿润2 块样地, 并对其中生长的胡杨(Populus euphratica)叶片的呼吸作用特性进行了观测。结果表明, 干旱样地胡杨叶片的相对含水量(RWC)和单位质量氮含量(Nmass)显著低于湿润样地, 但叶片单位面积氮含量(Narea)和比叶面积(SLA)差异不显著。叶片暗呼吸速率(Rn)和光下线粒体呼吸速率(RL)均随地下水埋深的降低而升高, 其中地下水埋深度的变化对RL 影响更加明显。光对叶片暗呼吸速率具有抑制作用, 其抑制程度在11%-57%变化。光对暗呼吸的抑制程度同Rubisco 酶的最大氧化速率(vo)呈负相关。湿润样地胡杨叶片光合速率(A)同光下线粒体呼吸速率的比值(A/RL)显著高于A 同Rn 的比值(A/Rn), 而在干旱样地2 者间差异不显著。胡杨叶片呼吸速率和光对暗呼吸抑制作用对地下水埋深的响应可能同细胞抗旱对能量和物质需求的增加有关。此外, 叶片A/R 比值不仅受到光照条件的影响, 同时受到水分条件的调控。     

增温对青藏高原高寒草原生态系统碳交换的影响

碳交换是影响草地生态系统碳汇功能的关键过程,对气候变暖极为敏感。青藏高原分布着大面积的高寒草原,其碳汇功能对气候变暖的响应对区域碳循环过程具有重要的影响。为探究高寒草原生态系统碳交换过程对增温的响应,2012—2014年,在青藏高原班戈县进行了模拟增温对高寒草原生态系统碳交换过程影响的研究。结果表明,增温对高寒草原碳交换各组分的影响存在年际差异,但总体上对碳交换存在负面影响。3年平均结果显示,增温显著降低了高寒草原地上生物量、总生态系统生产力(GEP)、生态系统呼吸(ER)和净生态系统碳交换量(NEE)(P0.05),平均降幅分别为15.1%、36.8%、19.2%和51.5%。增温条件下3年平均土壤呼吸(SR)较对照无显著变化(P0.05),但2013年增温显著降低了SR(P0.05),降幅达18.1%。增温对SR与ER的比值具有一定的促进作用,最高增幅达到40.0%。GEP、ER、SR和NEE与土壤温度和土壤水分无显著相关(P0.05),而GEP、ER和NEE与空气温度呈显著的负相关关系(P0.05)。增温引起的干旱胁迫以及地上生物量降低是导致高寒草原NEE降低的主要原因。研究表明,全球变暖会一定程度降低青藏高原高寒草原的碳汇功能。     

大气CO2浓度倍增对植物暗呼吸的影响

以长期生长于350和700μmolCO_2·mol~(-1)空气的开顶式培养室的杜仲(Eucommia ulmoides Oliv.)、紫花苜蓿(Medicago sativa L.)、玉米(Zea mays L.)等10种植物的离体成熟叶片或整株为材料,研究不同测定温度(15～35℃)下,CO_2浓度倍增对植物暗呼吸的影响。结果表明:在较低温度(15℃、20℃)下,CO_2浓度倍增对植物暗呼吸没有显著效应,在较高温度(30℃、35℃)下多数被测植物的暗呼吸显著增强。讨论了实验所得结果在未来全球气候变化中的可能的意义。     

红松和紫椴叶片暗呼吸及其光抑制性在幼、成树间的差异

叶片暗呼吸是森林碳循环的重要组分,深入分析幼、成树的叶片暗呼吸及其光抑制性的差异,对生态系统总生产力(GPP)的准确估算具有重要意义.本研究以长白山阔叶红松林主要树种(红松和紫椴)的幼树和成树为研究对象,分别测算不同光照下叶片暗呼吸与无光暗呼吸,比较叶片暗呼吸及其光抑制性在幼、成树间的差异,结合幼、成树叶片生理生态参数的对比,对幼、成树叶片暗呼吸及其光抑制性差异的原因进行探讨.结果表明: 两个树种幼树叶片光下暗呼吸的值高于成树,在生长季(6—9月),幼树的值比成树高6.8%～39.6%;两个树种幼树叶片暗呼吸光抑制程度低于成树,幼树叶片暗呼吸光抑制性的值比成树低2.5%～14.1%;红松幼、成树间叶片暗呼吸光抑制性的差异总体高于紫椴幼、成树间叶片暗呼吸光抑制性的差异,差值最高可达18.6%;幼树中较高的光下暗呼吸值和较低的光抑制程度可能与最大净光合速率、比叶面积、气孔导度的变化有关,与叶片氮含量的变化无关.     

温带4种针叶树种春、秋季节树干维持呼吸的日动态

揭示树干维持呼吸(RM)的时间变化特征及其调控因子有助于理解树木碳代谢过程及其对环境变化的响应和构建森林碳循环机理模型。采用红外气体分析法原位测定东北东部山区4个针叶树种(红松、红皮云杉、樟子松和兴安落叶松)的春、秋季节RM日动态及其影响因子。结果表明:秋季和春季4个树种RM日变化多随树干温度(TW)而变化,但RM峰值大小和出现时间以及日变化幅度因树种和季节而异。TW解释了RM(除春季樟子松外)变异性的50%以上,但RM对TW响应滞后1.5 h(春季樟子松为3 h)。将RM标准化到TW为10℃(R10)时发现,秋季R10波动在0.54μmol CO2m-2s-1(兴安落叶松)—0.78μmol CO2m-2s-1(红皮云杉)之间,而春季R10则波动在0.87μmol CO2m-2s-1(红松)—1.10μmol CO2m-2s-1(樟子松)之间,前者平均低于后者约40%。然而,各树种秋季和春季RM的Q10值差异不显著(P0.05),波动在1.52(樟子松)—1.82(红皮云杉)之间。秋季和春季所有树种的R10与树木胸径(DBH)之间均呈显著的正相关关系(P0.05),而Q10与DBH则多呈负相关关系(P0.05),表明DBH可作为估测这些针叶树种RM的参数之一。     

高寒矮嵩草草甸冬季CO2释放特征

冬季碳排放在高寒草地年内碳平衡中占有重要位置。为探讨高寒草地冬季碳排放特征及温度敏感性,于2003-2005年在中国科学院海北高寒草甸生态系统研究站,利用密闭箱-气相色谱法连续观测了高寒矮嵩草草甸2个冬季的生态系统、土壤呼吸通量特征。结果表明:1)高寒矮嵩草草甸冬季生态系统呼吸、土壤呼吸均具有明显的日变化和季节变化规律,温度是其主要的控制因子,能够解释44%以上的呼吸速率变异。2)冬季生态系统呼吸与土壤呼吸速率在统计上没有显著差异,土壤呼吸占生态系统呼吸的比例高达85%以上。3)2003-2004年冬季生态系统呼吸、土壤呼吸的Q₁₀值分别为1.53,1.38;2004-2005年冬季生态系统呼吸与土壤呼吸的Q₁₀值为1.86,1.68,2个冬季生态系统呼吸的Q₁₀值均高于土壤呼吸。4)未发现高寒矮嵩草草甸冷冬年份的Q₁₀值高于暖冬年份以及冬季的Q₁₀值高于生长季。     

亚热带六种常见榕属植物日间呼吸特征

植物呼吸作用是生态系统碳循环的一个关键过程。陆地植被每年通过光合作用固定的碳有一半被呼吸作用释放,而释放的这部分又有大约一半来自叶片呼吸。植物不仅在夜晚进行呼吸作用(R_d, Dark respiration),白天也在呼吸,即日间呼吸(R_L, Light respiration)。有研究表明光照条件下植物叶片呼吸低于黑暗环境下叶片呼吸速率,通常受到的光照抑制程度为30%—40%左右。但在实际研究中,经常忽视植物叶片日间呼吸的光抑制,进而大幅高估生态系统呼吸,从而使得生态系统总初级生产力(Gross primary productivity, GPP)被高估,因此有必要对植物的日间呼吸特性进行研究。以中国南方亚热带六种常见榕属植物(垂叶榕、高山榕、黄金榕、菩提树、细叶榕、小叶榕)为研究对象,测量其叶片的日间呼吸速率(R_L, Kok方法)、暗呼吸速率(R_d)、比叶重(LMA)、叶片氮含量(N)、磷含量(P)、最大光合速率(A)、1800及80μmol m^-2 s^-1     

华北平原玉米叶片光合及呼吸过程对实验增温的适应性

利用典型农田生态系统的原位实验增温平台,探讨我国华北平原重要农作物玉米叶片光合及呼吸过程对实验增温的适应性,并深入分析其产生适应性的原因和机理。研究结果显示,实验增温使玉米叶片净光合速率(A_n)显著升高(P0.001),同时增温也导致A_n的最适温度(T_(opt))升高1.56℃;相似地,实验增温也同样导致了光合作用过程中最大电子传递速率(J_(max))显著增加(P0.001),并且其最适温度(T_(opt))升高了1.45℃,但并没有对最大羧化反应速率(V_(cmax))及其温度敏感性(Q_(10))产生显著的影响(P0.05)。然而,实验增温却显著降低了玉米叶片的暗呼吸速率(R_d)及其Q_(10)值(P0.05)。另外,研究结果还显示实验增温没有对R_d/A_g和J_(max)/V_(cmax)产生显著的影响(P0.05)。此外,尽管实验增温显著提高了玉米叶片的蒸腾速率(T_r),但却并没有显著改变叶片的气孔导度(G_s)及水分利用效率(WUE)。研究结果表明,玉米可以通过调控叶片光合及呼吸等关键生理过程的最适温度对增温产生一定的适应性。然而,尽管玉米能够在叶片尺度上做出调整来适应增温环境,但这种适应能力却十分有限,以至于未来气候变暖仍可能会对华北平原玉米的生长发育过程和粮食产量造成一定的影响。     

生境不完全重叠的两种鲤科鱼类耐低氧及运动能力比较

以乌江流域亲缘关系近,但分布并不完全重叠的马口鱼(Opsariichthys bidens)和宽鳍鱲(Zacco platypus)作为实验对象,分别考察这两种实验鱼的低氧耐受及游泳运动能力。将野外采回的实验鱼置于(25±1)℃条件下,分别测定两种鱼的临界氧分压(Pcrit)、水面呼吸(ASR)、失去平衡点(LOE)以及在不同溶氧水平(8.0、4.0和2.0 mg/L)下的临界游泳速度(Ucrit)和活跃耗氧率(MO2active)。研究发现:马口鱼的Pcrit(2.44±0.20)mg/L显著高于宽鳍鱲(1.86±0.10)mg/L(P=0.031)。但马口鱼的50%ASR(1.23±0.16)mg/L显著低于宽鳍鱲(1.97±0.11)mg/L(P=0.023);马口鱼的50%LOE(0.84±0.01)mg/L同样显著低于宽鳍鱲(0.97±0.02)mg/L(P=0.004)。宽鳍鱲在8.0和4.0 mg/L下的游泳能力显著高于马口鱼,然而马口鱼和宽鳍鱲的Ucrit均随测定溶氧水平的下降而显著降低(P0.01);宽鳍鱲和马口鱼的运动耗氧率均随水流速度的增加而呈现指数增加,但随测定溶氧水平的降低运动耗氧曲线变得相对平缓,尤其是在溶氧为4.0 mg/L时马口鱼的运动耗氧曲线与宽鳍鱲相比越发平缓;两种实验鱼的MO2active随溶氧水平下降的变化趋势与Ucrit相似(P0.001)。结果表明:两种野外生存的实验鱼不仅在低氧耐受能力方面存在显著差异,而且两者的游泳运动能力也有不同表现,这很有可能与其遗传特征、生存环境及生态习性相关。     

Soil microbial activities and heavy metal mobility in long-term contaminated soils after addition of EDTA and EDDS

A 40-day incubation experiment was carried out in order to evaluate the microbial activities and heavy metal availability in long-term contaminated arable and grassland soils after addition of EDTA (ethylenediaminetetraacetic acid) or EDDS ([S,S]-ethylenediaminedisuccinic acid). Soils with similar contamination of heavy metal from the vicinity of a lead smelter were used in the experiment. The soil microbial carbon (C_mic) decreased significantly after addition of EDTA in the arable soil (CM1); lesser effects were observed in the grassland soil (CM2). Addition of EDDS caused a decrease of C_mic during the first 10 days of incubation. In the later phases of the experiment, C_mic increased, and even exceeded the amounts found in the control soils. Respiratory activities and metabolic quotients (qCO₂) increased after the addition of the chelating agents into the soils. Higher respiratory activities and qCO₂ were observed in the EDTA-treated soils. The readily available heavy metal fractions were extracted with NH₄NO₃ solution. Readily mobilizable heavy metal fractions of Cd, Pb, Zn, and (in part) Cu increased during the first 3-10 days of incubation in the presence of EDTA. The addition of EDDS particularly increased concentrations of available Cu. Significant correlations between NH₄NO₃-extractable metals, soil respiratory activities, and qCO₂ were found in both soil treatments with EDTA and EDDS. This indicates that enhanced metal mobility seriously affects the microbial processes in experimental soils. In addition, the relationships between NH₄NO₃-extractable Cd, Cu, and the microbial biomass were found in the CM1 soil amended with EDTA.     

Glucosinolates in seeds of Neslia paniculata

The glucosinolate fraction in seeds of Neslia paniculata (L.) Desv., subsp. thracica (Velen.) Bornmüller, (Cruciferae), has been isolated and subjected to enzymic hydrolysis. The isothiocyanates thus produced have been separated and identified as cheirolin (II), (R)-9-methylsulfinylnonyl (III), (R)-10-methylsulfinyldecyl (IV), and (R)-11-methylsulfinylundecyl isothiocyanate (V), the last of which has not been previously encountered as a natural compound.     

神农架海拔梯度上4种典型森林的土壤呼吸组分及其对温度的敏感性

量化森林土壤呼吸及其组分对温度的响应对准确评估未来气候变化背景下陆地生态系统的碳平衡极其重要。该文通过对神农架海拔梯度上常绿阔叶林、常绿落叶阔叶混交林、落叶阔叶林以及亚高山针叶林4种典型森林土壤呼吸的研究发现: 4种森林类型的年平均土壤呼吸速率和年平均异养呼吸速率分别为1.63、1.79、1.74、1.35 μmol CO₂·m^-2·s^-1和1.13、1.12、1.12、0.80 μmol CO₂·m^-2·s^-1。该地区的土壤呼吸及其组分呈现出明显的季节动态, 夏季最高, 冬季最低。4种森林类型中, 阔叶林的土壤呼吸显著高于针叶林, 但阔叶林之间的土壤呼吸差异不显著。土壤温度是影响土壤呼吸及其组分的主要因素, 二者呈显著的指数关系; 土壤含水量与土壤呼吸之间没有显著的相关关系。4种典型森林土壤呼吸的Q₁₀值分别为2.38、2.68、2.99和4.24, 随海拔的升高土壤呼吸对温度的敏感性增强, Q₁₀值随海拔的升高而增加。     

Semi-stochastic models for Salmonella infection within finishing pig units in the UK

A multi-group semi-stochastic model is formulated to describe Salmonella   dynamics on a pig herd within the UK and assess whether farm structure has any effect on the dynamics. The models include both direct transmission and indirect (via free-living infectious units in the environment and airborne infection). The basic reproduction number _R0$R_0$ is also investigated. The models estimate approximately 24.6% and 25.4% of pigs at slaughter weight will be infected with Salmonella within a slatted-floored and solid-floored unit respectively, which corresponds to values found in previous abattoir and farm studies, suggesting that the model has reasonable validity. Analysis of the models identified the shedding rate to be of particular importance in the control of Salmonella   spread, a finding also evident in an increase in the _R0$R_0$ value.     

Comparing methods to quantify experimental transmission of infectious agents

Transmission of an infectious agent can be quantified from experimental data using the transient-state (TS) algorithm. The TS algorithm is based on the stochastic SIR model and provides a time-dependent probability distribution over the number of infected individuals during an epidemic, with no need for the experiment to end in final-size (e.g., where no more infections can occur). Because of numerical limitations, the application of the TS algorithm is limited to populations with only a few individuals. We investigated the error of using the easily applicable, time-independent final-size (FS) algorithm knowing that the FS situation was not reached. We conclude that the methods based on the FS algorithm: (i) underestimate R(0), (ii) are liberal when testing H(0):R(0)1 against H(1):R(0)<1, (iii) are conservative when testing H(0):R(0)1 against H(1):R(0)>1, and (iv) are conservative when testing H(0):R(control)=R(treatment) against H(1):R(control)>R(treatment). Furthermore, a new method is presented to find a difference in transmission between two treatment groups (MaxDiff test). The MaxDiff test is compared to tests based on FS and TS algorithms. The TS test and the MaxDiff test were most powerful (approximately equally powerful) in finding a difference, whereas the FS test was less powerful (especially, when both R(control) and R(treatment) are >1).     

A comparison of low temperature tolerance traits between closely related aphids from the tropics, temperate zone, and Arctic

The survival of aphids exposed to low temperatures is strongly influenced by their ability to move within and between plants and to survive exposure to potentially lethal low temperatures. Little is known about the physiological and behavioural limitations on aphid movement at low temperatures or how they may relate to lethal temperature thresholds. These questions are addressed here through an analysis of the thermal ecology of three closely related aphid species: Myzus persicae, a ubiquitous temperate zone pest, Myzus polaris, an arctic species, and Myzus ornatus, a sub-tropical species. Lower lethal temperatures (LLT₅₀) of aphids reared at 15 °C were similar for M. persicae and M. polaris (range: −12.7 to −13.9 °C), but significantly higher for M. ornatus (−6.6 °C). The temperature thresholds for activity and chill coma increased with rearing temperature (10, 15, 20, and 25 °C) for all clones. For M. polaris and M. ornatus the slopes of these relationships were approximately parallel; by contrast, for M. persicae the difference in slopes meant that the difference between the temperatures at which aphids cease walking and enter coma increased by approximately 0.5 °C per 1 °C increase in rearing temperature. The data suggest that all three species have the potential to increase population sizes and expand their ranges if low temperature limitation is relaxed.     

A [P]NAD-based method to identify and quantitate long residence time enoyl-acyl carrier protein reductase inhibitors

The classical methods for quantifying drug–target residence time (t_R) use loss or regain of enzyme activity in progress curve kinetic assays. However, such methods become imprecise at very long residence times, mitigating the use of alternative strategies. Using the NAD(P)H-dependent FabI enoyl-acyl carrier protein (enoyl-ACP) reductase as a model system, we developed a Penefsky column-based method for direct measurement of t_R, where the off-rate of the drug was determined with radiolabeled [adenylate-³²P]NAD(P⁺) cofactor. In total, 23 FabI inhibitors were analyzed, and a mathematical model was used to estimate limits to the t_R values of each inhibitor based on percentage drug–target complex recovery following gel filtration. In general, this method showed good agreement with the classical steady-state kinetic methods for compounds with t_R values of 10 to 100 min. In addition, we were able to identify seven long t_R inhibitors (100–1500 min) and to accurately determine their t_R values. The method was then used to measure t_R as a function of temperature, an analysis not previously possible using the standard kinetic approach due to decreased NAD(P)H stability at elevated temperatures. In general, a 4-fold difference in t_R was observed when the temperature was increased from 25 to 37 °C.     

新疆杨边材贮存水分对单株液流通量的影响

树木体内贮存水分量的大小及其参与液流循环的程度被认为是树木抵御干旱胁迫的重要机制之一.以我国北方广泛适生的新疆杨为研究对象,利用热扩散技术(TDP)分别监测了树冠基部、树杆基部处液流通量,并结合同步气象观测,分析了树杆不同高度处液流通量与大气蒸发潜力(ET0)间的关系,发现:冠基部的液流通量(Qu)是估算新疆杨单株液流通量可靠的指标,试验期间(6-9月)典型晴天日Qu日平均值为(7.61±0.65) L/d,比云天、阴天分别高0.41和2.71 L/d.新疆杨杆基部液流通量(Qd)在不同季节变化较大,在前3个月的典型晴天日,Qd小于Qu,树木处于失水过程,但在8月下旬连续多日降雨后迅速增加而反超,新疆杨能通过这种方式缓解季节间的水分亏缺,维持水分的总体平衡.在8月下旬连续多日降雨、土壤水分得到有效改善后,Qd占全天液流总量的比例由先前的31.98％-35.36％下降到6.72％-7.99％,夜间液流成为补偿与缓解日间水分亏缺的重要方式.在整个生长季内,新疆杨体内经历了水分补充(6月)—失水(7、8月份)—再补充(9月)的过程.ET0是评价液流环境驱动效应可靠的综合变量,建立基于Logistic方程的模型可通过ET0较好的估算Qu,估算新疆杨Qu的上限约为7.82L/d.分析显示当ET0超过5mm/d时,Qu、Qd均不再随ET0的增加而增加或有下降,显示了树木主动保护、抵御干旱的策略.8月下旬连续降雨使得新疆杨蒸腾量占大气蒸发潜力的比例(T/ET0)由先前的0.32增加到之后的0.47,可以看出土壤水分改善对液流量的贡献较大.     

Notions of synergy for combinations of interventions against infectious diseases in heterogeneously mixing populations

In public health programmes interventions are frequently combined with hoped for ‘synergies’ [22]. However, there is not yet a precise definition for synergy between interventions that captures the idea that there is added benefit at the population-level in using them together. To explore the synergy between interventions in the context of endemic disease, we consider a general model of infection spread in a heterogeneously mixing population. We consider interventions which may alter individuals’ infectiousness, susceptibility, profile of infectiousness through time and survival while infected. Allowing general patterns of overlap and targeting in those receiving the interventions, we show how to compute changes to epidemiological indices such as R₀, and introduce a simple technique for calculating equilibrium prevalences and incidences via an iterated map. We argue for a particular definition of synergy and investigate its behaviour, both analytically and numerically, concluding that it is easiest to achieve synergy between interventions which perform poorly in isolation; implementation strategies that minimize the overlap of different interventions in the population tend to achieve more synergy; and that in populations with heterogeneous risk, interventions that are redundant when universally targeted can regain substantial synergy when applied in a targeted manner.