Monitoring conidial density of <Emphasis Type="Italic">Monilinia fructigena</Emphasis> in the air in relation to brown rot development in integrated and organic apple orchards

In a three-year Hungarian study, conidial density of Monilinia fructigena in the air determined from mid-May until harvest was related to brown rot disease progress in integrated and organic apple orchards. Conidia of M. fructigena were first trapped in late May in both orchards in all years. Number of conidial density greatly increased after the appearance of first infected fruit, from early July in the organic and from early August in the integrated orchard. Conidial number continuously increased until harvest in both orchards. Final brown rot incidence reached 4.3–6.6% and 19.8–24.5% in the integrated and organic orchards, respectively. Disease incidence showed a significant relationship with corresponding cumulative numbers of trapped conidia both in integrated and organic orchards, and was described by separate three-parameter Gompertz functions for the two orchards. Time series analyses, using autoregressive integrated moving average (ARIMA) models, revealed that the temporal patterns of the number of airborne conidia was similar in all years in both integrated and organic orchards. Conidia caught over a 24-h period showed distinct diurnal periodicity, with peak spore density occurring in the afternoon between 13.00 and 18.00. Percent viability of M. fructigena conidia ranged from 48.8 to 70.1% with lower viability in dry compared to wet days in both orchards and all years. Temperature and relative humidity correlated best with mean hourly conidial catches in both integrated and organic apple orchards in each year. Correlations between aerial spore density and wind speed were significant only in the organic orchard over the 3-year period. Mean hourly rainfall was negatively but poorly correlated with mean hourly conidial catches. Results were compared and discussed with previous observations.     

Conidial dispersal by Alternaria brassicicola on Chinese cabbage (Brassica pekinensis) in the field and under simulated conditions

This study investigated conidial dispersal in the field, and effects of simulated wind and rain on the dispersal of A. brassicicola on Chinese cabbage ( Brassica pekinensis ). Spores were sampled using a Burkard volumetric spore sampler and rotorod samplers in a Chinese cabbage crop. Disease incidence in the field was well fitted by a Gompertz curve with an adjusted r ² of >0·99. Conidia of A. brassicicola were trapped in the field throughout the growing season. Peaks of high spore concentrations were usually associated with dry days, shortly after rain, high temperature or high wind speed. Diurnal periodicity of spore dispersal showed a peak of conidia trapped around 10·00 h. The number of conidia trapped at a height of 25 cm above ground level was greater than that at 50, 75 and 100 cm. Conidial dispersal was also studied under simulated conditions in a wind tunnel and a rain simulator. Generalized linear models were used to model these data. The number of conidia caught increased significantly at higher wind speeds and at higher rain intensities. Under simulated wind conditions, the number of conidia dispersed from source plants with wet leaves was only 22% of that for plants with dry leaves. Linear relationships were found between the number of conidia caught and the degree of infection of trap plants.     

Effects of humidity, leaf wetness, temperature and light on conidial production by Phaeoisariopsis personata on groundnut

Controlled-environment studies of conidial production by Phaeoisariopsis personata on groundnut are described. With constant relative humidity (RH), conidia were only produced above a threshold (94·5% RH) and there was a linear increase between 94·5% RH and 100% RH. Conidial production was less with continuous leaf wetness (resembling heavy dew) than with continuous 98–99% RH, but it was similar with intermittent leaf wetness and intermittent 98–99% RH (8 h at 70% RH each day). With alternate high (≥97% RH) and low humidity, daily conidial production depended both on the duration of high RH and on the low RH value. With 99% RH at night (12 h), night-time conidial production decreased with the previous daytime RH. After conidial production had started, small numbers of conidia were produced even when the RH was well below the threshold (94·5%). Conidia were produced in continuous light when the photon flux density was 2 μmol/m²/s, but production was completely inhibited with 60 μmol/m²/s. With constant RH, more conidia were produced with a 12 h photoperiod than in continuous darkness. However, more than 75% of the conidia were produced in the dark. With continuous darkness, more conidia were produced during the night (18.00–06.00 h) than during the day, but this biological rhythm was overcome with a (light-night)/(dark-day) regime. With constant 98–99% RH there was a linear increase in conidial production with temperature between 10 and 28°C, and virtually no conidia were produced at 33°C. The daily production of conidia increased with time for 2 to 6 days, depending on the treatment.     

Relationships between Weather Variables, Airborne Spore Concentrations and Severity of Leaf Blight of Garlic Caused by Stemphylium vesicarium in Spain

Ascospores and conidia released into the air were recorded around plots on which garlic debris infected by Stemphylium vesicarium were fixed onto the soil surface. Symptoms in garlic trap plots located in the vicinity of infected debris, started in March and developed during April–May to reach disease incidence close to 100%, final disease severity values being lower in 1993 and 1995 than in 1994 and 1996. Whereas daily concentrations of ascospores were rather erratic, with 30% of captures between 0 and 6 h, conidia showed a daily periodicity with highest concentrations between 12 and 18 h, with a pronounced peak between 14 and 16 h, and lowest values at night. Ascospore release occurred mainly in February and March. It coincided with rainfall periods, 14 h with vapour pressure deficit 5 mb and solar radiation <145 W m^–2 on the current day of the capture. In contrast, greatest captures of conidia started in late April and were prevalent in May, and were associated with rainfall in days previous to the capture in which rather high temperature occurred and solar radiation was 109–345 W m^–2. Among the weather variables considered, rainfall appeared directly related to the aerial concentration of ascospores and conidia. The role of relative humidity seemed essential when rainfall did not occur. There was a relationship between conidia concentration in the air and number of hours with temperature in the range 12–21 °C. Ascospore production was not essential for infections to take place, since primary infection from conidia may occur and disease can develop from them readily.     

Relationship between Concentrations of Botrytis Cinerea Conidia in Air, Environmental Conditions, and the Incidence of Grey Mould in Strawberry Flowers and Fruits

Atmospheric concentrations of Botrytis cinerea conidia were monitored for two seasons in a strawberry crop in Moguer (Huelva, southwestern Spain). Concentrations of conidia were estimated using a Burkard volumetric spore sampler. A diurnal pattern of conidial release was observed. Airborne conidial concentration was significantly and positively correlated with the average solar radiation and mean temperature, and negatively with rainfall and relative humidity. Among the weather variables considered, solar radiation showed the most consistent results in the regression analysis, explaining over 40% of airborne conidial concentration variability. Correlation between Botrytis fruit rot incidence and accumulated number of conidia over seven days was significant and positive. Two regression models containing three variables explained over 62 and 52% of the fruit rot incidence variability. A positive but non-significant correlation was established between B. cinerea incidence in flowers and airborne conidial concentration. It was not possible to fit a consistent regression model to relate flower infection incidence to conidial concentration or weather variables.     

Effect of water on germination of Plasmopara viticola oospores

The effect of moisture in grape leaf litter holding overwintering Plasmopara viticola oospores was investigated. Oospores were incubated under different regimes of water activity ( a_W 0·991 to 0·123) for 2 to 15 days and their ability to germinate and cause infection was determined using a sensitive leaf disk assay. Reduction of a_W caused a significant shift in the infection dynamics, with maximum effect when a_W  ≤ 0·56. Dynamics of a_W in the leaf litter under natural conditions were estimated from moisture data using a Chen-Clayton equation. Daily patterns of leaf litter moisture (M in % weight) were determined in non rainy periods between mid February and mid June, while the Chen-Clayton equation was calculated using data of a_W and M measured in both sorption and desorption conditions, at different temperatures. Water activity was highest at 08·00 hours, decreased progressively until 14·00 hours, and then increased. Water activity was favourable for oospores to develop in about 25% of the measurements, all made between 18·00 hours and 08·00 hours. A close relationship was found between vapour pressure deficit (VPD in hPa) and a_W of the leaf litter, so that when VPD is lower than 2·13 hPa there is sufficient water for oospores to develop. Results showed that leaf litter moisture due to water from the atmosphere makes oospore development possible during non rainy periods.     

北京地区日光温室草莓灰霉病发生 动态监测和影响因素分析

 草莓灰霉病是由灰葡萄孢(Botrytis cinerea Pers.)引起的一种真菌病害,可造成草莓烂果,严重影响草莓产量和采后保存。为了探明日光温室草莓灰霉病的发生流行规律,在2013—2014年生长季和2014—2015年生长季,对北京地区草莓日光温室空气中灰葡萄孢分生孢子数量、草莓花瓣带菌率和灰霉病病果数进行了动态监测和调查,同时对日光温室中气象因子进行了系统监测和记录。结果表明,在草莓日光温室中,利用孢子捕捉器捕获的灰葡萄孢分生孢子数量在一天中主要集中在5:00-18:00,以11:00-14:00数量最大。在一天中每小时捕获的分生孢子数量与温度和光照强度呈极显著正相关(P≤0.01),与相对湿度呈极显著负相关(P≤0.01)。新增草莓灰霉病病果数与7 d前当天捕获的分生孢子数量呈极显著正相关(r=0.872,P≤0.01),与7 d前当天的新鲜花瓣带菌率亦呈极显著正相关(r=0.807,P≤0.01),这为利用捕获的分生孢子数量和新鲜花瓣的带菌率预测7 d后草莓灰霉病的发生情况提供了重要参考。本研究结果有助于了解日光温室中草莓灰霉病的发生规律和影响因素,为该病害的防控和预测测报提供了依据。     

生长季马铃薯植株冠层空气中Alternaria solani分生孢子飞散动态及其影响因素

 对生长季马铃薯冠层空气中Aternaria solani (Ell. & G. Martin ) 分生孢子的数量和马铃薯早疫病发生程度进行了系统调查,并对空气中分生孢子数量与早疫病发生程度之间的相关性以及影响空气中分生孢子数量的因素进行了分析。结果表明:在马铃薯出苗至收获(6~8月份)期间,当病情指数低于4时两者之间呈显著线性相关关系,随着病情指数的增高二者的线性相关性降低。天气条件对空气中A. solani 分生孢子数量的影响明显,通常降雨2 d后空气中孢子的数量会有一个高峰,日平均气温(地上2 m)低于18℃的条件下,空气中孢子数量会显著减少。一天中6∶00 am至16∶00 pm空气中A. solani分生孢子数量较多,17∶00 pm至次日5∶00 am孢子数量较少,分别占全天总量的 60.95%和 39.05%,各小时孢子飞散量占全天比例与空气相对湿度呈显著负相关,而与空气温度呈显著正相关。     

Dynamics in concentrations of <Emphasis Type="Italic">Blumeria graminis</Emphasis> f. sp <Emphasis Type="Italic">tritici</Emphasis> conidia and its relationship to local weather conditions and disease index in wheat

Experiments were conducted for 3 seasons, 2007–2008, 2008–2009 and 2009–2010 in a wheat field planted with a cultivar susceptible to powdery mildew in Langfang City, Hebei Province, China. Plants were inoculated with Blumeria graminis f. sp. tritici (Bgt) and conidia of Bgt in the air were trapped using volumetric spore samplers. Disease severity was recorded weekly. The relationships between airborne conidial concentrations and meteorological factors, as well as disease index were analyzed. Conidia were first detected about 20 days after inoculation in all three seasons, and then increased gradually with time. The highest conidial concentrations in the air were observed in mid-May 2008 and 2009 and late May 2010 at growth stage (GS) 10.5.4. The concentrations of Bgt conidia after inoculation (GS 5) to milky ripe (GS 11.1) in the air were positively correlated with temperature, solar radiation, and negatively with relative humidity and vapor pressure deficit (VPD). Prediction models of Bgt conidial concentrations in the air based on meteorological factors were constructed using multiple regression analysis. Time series analysis, using autoregressive integrated moving average (ARIMA) (p, d, q) models, showed that each of the three season’s data can be fitted with simple ARIMA (1, 0, 0) models. Conidial concentrations within the canopy were significantly higher than those above the canopy (P < 0.01). The weekly-accumulated mean hourly conidia per cubic metre of air significantly (P < 0.01) correlated with disease index in all three seasons.     

Mummified fruit as a source of inoculum and disease dynamics of olive anthracnose caused by Colletotrichum spp

Anthracnose, caused by Colletotrichum spp., is a destructive disease of olive fruit worldwide. The objective of this study was to investigate the influence of agronomical and weather factors on inoculum production using detached olive fruit and on the development of epidemics in the field. The pathogen produced very large numbers of conidia on rotted (>1.87 × 10(8) conidia/fruit) or mummified (>2.16 × 10(4) conidia/fruit) fruit under optimal conditions. On mummified fruit, conidial production was highest on mummies incubated at 20 to 25°C and 96 h of wetness. Repeated washings of mummies reduced conidial production until it was very low after five washings. When mummies were placed in the tree canopy, conidial production was not reduced after 6 months (May to October); but, when they were held on the soil or buried in the soil, conidial production comparatively decreased up to 10,000 times. Anthracnose epidemics on susceptible 'Hojiblanca' and 'Picudo' during three seasons (2005-08) were influenced by rainfall, temperature, and fruit ripening, and had three main phases: the latent period (May to October); the onset of the epidemic, which coincided with the beginning of fruit ripening (early November); and disease development, which was predicted by the Weibull model (November to March). No epidemics developed on the susceptible cultivars during the driest season (2007-08) or on the resistant 'Picual' olive during any of the three seasons. These results provide the basis for a forecasting system of olive anthracnose which could greatly improve the management of this disease.     

Sources and seasonal dynamics of inoculum for brown spot disease of pear

The dynamics of the production of Stemphylium vesicarium conidia and Pleospora allii ascospores from different inoculum sources on the ground were compared in a model system of a wildflower meadow mainly composed of yellow foxtail, creeping cinquefoil and white clover. The meadow was either inoculated (each October) or not inoculated with a virulent strain of S. vesicarium, and either covered or not covered with a litter of inoculated pear leaves. Spore traps positioned a few centimetres above the ground were exposed for 170 7-day periods between October 2003 and December 2006. Ascospores and conidia were trapped in 46 and 25% of samples, respectively. Ascospore numbers trapped from the pear leaf litter were about five times higher than those from the meadow, while conidial numbers were similar from the different inoculum sources. The ascosporic season was very long, with two main trapping periods: December–April, and August–October; the former was most important for the leaf litter, the latter for the meadow. The conidial season lasted from April to November, with 92% of conidia caught between July and September. The fungus persistently colonized the meadow: the meadow inoculated in early October 2003 produced spores until autumn 2006. The present work demonstrates that orchard ground is an important source of inoculum for brown spot of pear. Thus, it is important to reduce inoculum by managing the orchard ground all year long.     

Environmental Factors Affecting Production, Release, and Field Populations of Conidia of Alternaria alternata, the Cause of Brown Spot of Citrus

ABSTRACT Alternaria brown spot, caused by Alternaria alternata pv. citri, affects many tangerines and their hybrids, causing loss of immature leaves and fruit and reducing the marketability of the remaining fruit. Conidial production of A. alternata was greatest on mature leaves moistened and maintained at near 100% relative humidity (RH) for 24 h, whereas leaves that had been soaked or maintained at moderate RH produced few conidia. Conidial release from filter paper cultures and infected leaves was studied in a computer-controlled environmental chamber. Release of large numbers of conidia was triggered from both substrates by sudden drops in RH or by simulated rainfall events. Vibration induced release of low numbers of conidia, but red/infrared irradiation had no effect. In field studies from 1994 to 1996, air sampling with a 7-day recording volumetric spore trap indicated that conidia were present throughout the year with periodic large peaks. The number of conidia captured was not closely related to rainfall amounts or average wind speed, but was weakly related to the duration of leaf wetness. Likewise, disease severity on trap plants placed in the field weekly during 1995 to 1996 was not closely related to conidial numbers or rainfall amounts, but was weakly related to leaf wetness duration. Sufficient inoculum appears to be available to allow infection to occur throughout the year whenever susceptible host tissue and moisture are available.     

Temporal spore dispersal patterns of grapevine trunk pathogens in South Africa

Trunk disease pathogens of grapevines, viz. Phaeomoniella chlamydospora, Eutypa lata and several species in Botryosphaeriaceae, Phaeoacremonium and Phomopsis are known to infect fresh pruning wounds by means of air-borne inoculum released after rainfall or prolonged periods of high relative humidity. Recent surveys have demonstrated that most or all of these pathogens are present in climatically diverse grape growing regions of South Africa. However, the factors controlling spore dispersal of these pathogens in vineyards were largely unknown. To address this question, spore trapping was done in a Chenin Blanc vineyard in the Stellenbosch area, South Africa, for 14 weeks during the grapevine pruning period from June to mid-September of 2004 and 2005. Hourly recordings of weather data were done by a weather station in the row adjacent to the spore trap. Spores of E. lata and Phomopsis and species in Botryosphaeriaceae were trapped throughout the trapping periods of 2004 and 2005, with higher levels of trapped spores recorded in 2005. The spores of all three pathogens were trapped during or after periods of rainfall and/or high relative humidity. In neither of the 2 years were spores of Pa. chlamydospora or Phaeoacremonium spp. trapped. Results indicated that spore event incidence, as well as the amount of spores released during a spore event of above-mentioned pathogens, were governed by rainfall, relative humidity, temperature and wind speed prior to and during the spore events.     

Temporal patterns of airborne conidia of Podosphaera leucotricha, causal agent of apple powdery mildew

In each of the five years 1969 and 1971-1974 inclusive a volumetric spore trap was used in an apple orchard to monitor changes in the number of airborne conidia of Podosphaera leucotricha , the causal agent of apple powdery mildew. The number of trapped conidia varied greatly between years. Time-series analyses, using autoregressive integrated moving average (ARIMA) models, revealed that the temporal pattern of the number of airborne conidia was similar in all years, generally following a diurnal pattern with an afternoon peak. A strong correlation between consecutive hourly counts indicated that the number of trapped conidia depended on the strength of sporulating sources. Using the time-series transfer function (TF) method, it was shown that in each year the most important weather variables influencing the number of airborne conidia were vapour pressure deficit (VPD) and rainfall. Variation between years in the dynamic effects of these variables on conidium numbers was detected, and may reflect weather differences between years. Stepwise regression analysis was applied to the combined daily data for 1973 and 1974 using a subset of weather variables as independent variables, chosen on the basis of TF analysis. A resulting regression model accurately predicted the temporal pattern of conidium numbers (expressed as a percentage of the maximum daily number trapped in the same year) in both years. When this model was applied to the other three years there was good agreement between predicted and observed temporal patterns. Application of this regression model for practical disease control is discussed.     

Windborne spread of ergot disease (Claviceps africana) in sorghum A-lines in Zimbabwe

Ergot disease spread rapidly in Zimbabwe amongst replicated plots of male-sterile sorghum A-lines, from a group of centrally situated and precociously inoculated plants. Prominent secondary conidiation by the pathogen, Claviceps africana , on the surface of exuded honeydew provided airborne spores which were trapped in a Burkard continuous spore trap and showed diurnal peaks of concentration in air close to the primary source of inoculum. The rate of disease spread ( r =0·2; range 0·14–0·58) closely matched that recorded for other plant pathogens such as Phytophthora infestans and Puccinia graminis tritici , and it is concluded that the characteristic secondary conidia of C. africana were the principal epidemiological agents within the experimental area. Ergot spread by windborne secondary conidia has significant epidemiological and economic implications for sorghum hybrid breeding in southern Africa.     

Modelling the effect of cuticular crack surface area and inoculum density on the probability of nectarine fruit infection by Monilinia laxa

The effects of cuticular crack surface area and inoculum density on the infection of nectarine fruits by conidia of Monilinia laxa were studied using artificial inoculations with conidial suspensions and dry airborne conidia during the 2004 and 2005 seasons, respectively. Additionally, the effect of ambient humidity on fruit infection was evaluated in the 2005 experiment. An exploratory analysis indicated that (i) ambient humidity did not significantly explain the observed variability of data, but that (ii) the incidence of fruit infection increased both with increasing inoculum density and increasing surface area of cuticular cracks. The product of these two variables represented the inoculum dose in the cracks, and was used as a predictor of fruit infection in the model. Natural infection in the orchard was observed to increase throughout the season in both 2004 and 2005. The relationship between the probability of fruit infection by M. laxa and the artificially inoculated dose in the cuticular cracks was well described by a logistic regression model once natural inoculum density was taken into account (pseudo R² = 65%). This function could be helpful for estimating the risk of fruit infection at harvest based on fruit size and natural inoculum density.     

宁夏温室瓜菜白粉病菌鉴定及病害流行预测模型构建

为明确宁夏回族自治区温室瓜菜白粉病菌的分类地位,对采自该地区温室的南瓜、黄瓜和甜瓜上的白粉病菌基于ITS序列分析进行分子鉴定;利用孢子捕捉器对温室中甜瓜白粉病菌的孢子量进行监测,分析环境因子、孢子量和病情指数之间的关系,并采用逐步回归分析法构建温室甜瓜白粉病的流行预测模型。结果表明,基于ITS序列的分子鉴定结果,3种瓜菜白粉病的病原菌均为单囊壳白粉菌Podosphaera xanthii。发病期间,每日温室中甜瓜白粉病菌的孢子量在12:00—16:00时段最多,占24 h内总孢子量的34%~81%,20:00—08:00时段最少;白粉病菌孢子的释放与光照强度有关,相关系数为0.602。第t天的病情指数与标准累积温度、标准累积湿度、t-4 d前08:00—12:00时段的累积孢子量、第t-4天16:00—20:00时段的孢子量均具有显著的相关性,相关系数分别为0.935、0.938、0.956和0.921。以标准累积湿度和第t-4天16:00—20:00时段的孢子量为预测变量构建了温室甜瓜白粉病流行预测模型,决定系数为0.962,表明该模型具有较好的实际应用价值。     

温度对绿僵菌LA菌株孢子萌发、酶活力及毒力的影响

为明确温度对绿僵菌LA菌株生物学特性和毒力的影响,采用微室萌发、酶标仪测定A值以及体外涂抹法,对不同温度下LA菌株孢子萌发率、胞外蛋白酶、几丁质酶活力以及对东亚飞蝗的毒力进行了测定,结果表明:LA菌株在30℃下孢子萌发速率最快,对东亚飞蝗的致死速率也最快;38℃下培养,孢子萌发率开始受到限制,40℃下培养孢子的萌发率接近于0;高温处理转至适温培养,对孢子萌发速率产生一定影响,48℃下处理不同时间,孢子萌发速率、萌发率均显著降低,转至30℃下培养24h仅有10.6%～20.9%的孢子可萌发;LA胞外蛋白酶活力培养至第6天达到最高,为31.37U/mL,不同温度下活力测定表明37℃下胞外蛋白酶活力最高,几丁质酶在培养至第5天酶活力达到最高,为34.62U/mL,最高酶活温度为50℃。     

马铃薯早疫病菌分生孢子萌发条件的研究

为了明确马铃薯早疫病菌分生孢子萌发的条件,研究了温度、湿度、pH、光照、营养条件等对马铃薯早疫病菌分生孢子萌发的影响。结果表明,分生孢子在水滴中于30℃1 h即可萌发,8 h后达到萌发高峰,萌发率为98.4%;萌发的最适温度为30℃;相对湿度85%以上均可萌发,水滴中萌发率最高;pH6~10的条件利于分生孢子萌发;紫外线对分生孢子萌发有抑制作用;无机氮源中的铵态氮和尿素对孢子萌发有抑制作用,有机氮源中半胱氨酸和谷氨酸对孢子萌发有抑制作用;山梨糖不利于分生孢子萌发;光照对分生孢子萌发也有抑制作用;分生孢子致死温度是57℃。     

Estimating the germination dynamics of Plasmopara viticola oospores using hydro-thermal time

The effects of environmental conditions on the variability in germination dynamics of Plasmopara viticola oospores were studied from 1999 to 2003. The germination course was determined indirectly as the relative infection incidence (RII) occurring on grape leaf discs kept in contact with oospores sampled from a vineyard between March and July. The time elapsed between 1 January and the infection occurrence was expressed as physiological time, using four methods: (i) sums of daily temperatures > 8°C; (ii) hourly temperatures > 10°C; (iii) sums of hourly rates from a temperature-dependent function; or (iv) sums of these rates in hours with a rain or vapour pressure deficit ≤ 4·5 hPa (hydro-thermal time, HT). An equation of Gompertz in the form RII = exp[− a  · exp(− b  · HT)] produced an accurate fit for both separate years ( R ² = 0·97 to 0·99) and pooled data ( R ² = 0·89), as well as a good accuracy in cross-estimating new data ( r between observed and cross-estimated data were between 0·93 and 0·99, P  < 0·0001). It also accounted for a great part of the variability in oospore germination between years and both between and within sampling periods. Therefore, the equation of Gompertz (with a  = 15·9 ± 2·63 and b  = 0·653 ± 0·034) calculated over hydro-thermal time, a physiological time accounting for the effects of both temperature and moisture, produced a consistent modelling of the general relationships between the germination dynamics of a population of P. viticola oospores and weather conditions. It represents the relative density of the seasonal oospores that should have produced sporangia when they have experienced favourable conditions for germination.