Predicting germination response to temperature. III. Model validation under field-variable temperature conditions

BACKGROUND AND AIMS: Two previous papers in this series evaluated model fit of eight thermal-germination models parameterized from constant-temperature germination data. The previous studies determined that model formulations with the fewest shape assumptions provided the best estimates of both germination rate and germination time. The purpose of this latest study was to evaluate the accuracy and efficiency of these same models in predicting germination time and relative seedlot performance under field-variable temperature scenarios. METHODS: The seeds of four rangeland grass species were germinated under 104 variable-temperature treatments simulating six planting dates at three field sites in south-western Idaho. Measured and estimated germination times for all subpopulations were compared for all models, species and temperature treatments. KEY RESULTS: All models showed similar, and relatively high, predictive accuracy for field-temperature simulations except for the iterative-probit-optimization (IPO) model, which exhibited systematic errors as a function of subpopulation. Highest efficiency was obtained with the statistical-gridding (SG) model, which could be directly parameterized by measured subpopulation rate data. Relative seedlot response predicted by thermal time coefficients was somewhat different from that estimated from mean field-variable temperature response as a function of subpopulation. CONCLUSIONS: All germination response models tested performed relatively well in estimating field-variable temperature response. IPO caused systematic errors in predictions of germination time, and may have degraded the physiological relevance of resultant cardinal-temperature parameters. Comparative indices based on expected field performance may be more ecologically relevant than indices derived from a broader range of potential thermal conditions.     

Predicting germination response to temperature. I. Cardinal-temperature models and subpopulation-specific regression

BACKGROUND AND AIMS: The purpose of this study was to compare the relative accuracy of different thermal-germination models in predicting germination-time under constant-temperature conditions. Of specific interest was the assessment of shape assumptions associated with the cardinal-temperature germination model and probit distribution often used to distribute thermal coefficients among seed subpopulations. METHODS: The seeds of four rangeland grass species were germinated over the constant-temperature range of 3-38 degrees C and monitored for subpopulation variability in germination-rate response. Subpopulation-specific germination rate was estimated as a function of temperature and residual model error for three variations of the cardinal-temperature model, non-linear regression and piece-wise linear regression. The data were used to test relative model fit under alternative assumptions regarding model shape. KEY RESULTS: In general, optimal model fit was obtained by limiting model-shape assumptions. All models were relatively accurate in the sub-optimal temperature range except in the 3 degrees C treatment where predicted germination times were in error by as much as 70 d for the cardinal-temperature models. CONCLUSIONS: Germination model selection should be driven by research objectives. Cardinal-temperature models yield coefficients that can be directly compared for purposes of screening germplasm. Other model formulations, however, may be more accurate in predicting germination-time, especially at low temperatures where small errors in predicted rate can result in relatively large errors in germination time.     

ESS germination strategies in randomly varying environments. II. Reciprocal Yield-Law models

The study of ESS germination fractions in S. Ellner (1985, Theor. Pop. Biol. 27, 000-000) is applied and extended in the case of annuals obeying the Reciprocal Yield Law. The effects of parameter changes on the value of the ESS germination fraction are determined in some limiting cases by analyses of approximations. Numerical solutions of the ESS criterion are used to check the robustness of the conclusions and the accuracy of the approximations. In general, the ESS germination fraction decreases with an increased survivorship of buried seeds and with increased "variability" of seed yields. However, different measures of "variability" are appropriate in different circumstances. To avoid the possibility of conflicting predictions depending on the measure of variability, it is suggested that tests of the theory be limited to co-occurring species, and to variability due to climatic fluctuations. The ESS theory based on the Reciprocal Yield Law is compared with D. Cohen's (1966, J. Theor. Biol. 12, 119-129; 1968, J. Ecol. 56, 219-228) density-independent theory of "optimal" germination. The theories differ qualitatively and quantitatively regarding the influence of mean yield, seed survivorship, and the frequency of favorable years on the predicted germination fraction.     

温带半干旱地区一年生植物种子的萌发特性

在实验室条件下研究了中国温带半干旱地区科尔沁沙地的23种1年生植物的种子萌发特性（新采集种子、冷藏和干藏种子）.大籽蒿、虎尾草、冠芒草、沙蓬和地锦的新种子萌发率达90%左右,11种植物新种子萌发率均低于70%,说明这些植物的新种子具有或多或少的休眠属性.经过150d的冷干藏后,大籽蒿、虎尾草萌发率保持在90%以上,说明这两种植物完全没有休眠机制;冠芒草、沙蓬和地锦的种子萌发率下降较多,可能是储藏的环境条件导致的2次休眠现象;冷藏和干藏处理均能使绿珠藜、毛马唐、细叶益母草、雾冰藜、金狗尾草、苋菜、马齿苋、碱地肤和水稗草的种子在生长季开始时完成生理后熟,萌发率达到80%以上;干藏有利于促进毛马唐、细叶益母草、马齿苋和鹤虱的种子成熟,冷藏有利于促进绿珠藜和金狗尾草的种子成熟;黄蒿、灰绿藜、画眉草和烛台虫实在不同处理下的萌发率都比较低,说明种子内在生理休眠作用较强,具有减少种子一次性萌发数量的风险分摊策略.大多数1年生植物均能在较短时间内达到最终萌发率的90%,表现出迅速萌发的特性;黄蒿、灰绿藜、碱地肤和沙蓬种子则在不同处理中表现出延长萌发时间的策略来适应半干旱地区不确定的环境条件.最后,探讨了几种主要1年生植物的种子萌发对策与其对环境适应机制之间的关系.     

Seed germination and seedling physiology of Larix kaempferi and Pinus densiflora in seedbeds with charcoal and elevated CO2

We investigated the effect of ectomycorrhizal colonization, charcoal and CO₂ levels on the germination of seeds of Larix kaempferi and Pinus densiflora, and also their subsequent physiological activity and growth. The seeds were sown in brown forest soil or brown forest soil mixed with charcoal, at ambient CO₂ (360 μmol mol⁻¹) or elevated CO₂ (720 μmol mol⁻¹), with or without ectomycorrhiza. The proportions of both conifer seeds that germinated in forest soil mixed with charcoal were significantly greater than for seeds sown in forest soil grown at each CO₂ level (P < 0.05; t-test). However, the ectomycorrhizal colonization rate of each species grown in brown forest soil mixed with charcoal was significantly lower than in forest soil at each CO₂ treatment [CO₂] (P < 0.01; t-test). The phosphorus concentrations in needles of each seedling colonized with ectomycorrhiza and grown in forest soil were greater than in nonectomycorrhizal seedlings at each CO₂ level, especially for L. kaempferi seedlings (P < 0.05; t-test), but the concentrations in seedlings grown in brown forest soil mixed with charcoal were not increased at any CO₂ level. Moreover, the maximum net photosynthetic rate of each seedling for light and CO₂ saturation (P _max) increased when the seedlings were grown with ectomycorrhiza at 720 μmol mol⁻¹ [CO₂]. Ectomycorrhizal colonization led to an increase in the stem diameter of each species grown in each soil treatment at each CO₂ level. However, charcoal slowed the initial growth of both species of seedling, constraining ectomycorrhizal development. These results indicate that charcoal strongly assists seed germination and physiological activity.     

Proliferation of anchorage-dependent contact-inhibited cells. II: experimental results and validation of the theoretical models

This study establishes that the cellular automata models developed in an earlier article capture the essential features of the proliferation process for anchorage-dependent contact-inhibited cells. Model predictions are in excellent agreement with experimental data obtained with bovine pulmonary artery endothelial cells. The models are particularly suitable for predictive purposes since they have no adjustable parameters. All model parameters can be easily obtained from a priori measurements. Our studies also show that proliferation rates are very sensitive to the spatial distributions of seed cells. The adverse effects of seeding heterogeneities become more pronounced as a cell population approaches confluency and they should be accounted for in experimental studies attempting to assess the response of cells to external stimuli.     

Predicting germination response to temperature. II. Three-dimensional regression, statistical gridding and iterative-probit optimization using measured and interpolated-subpopulation data

BACKGROUND AND AIMS: Most current thermal-germination models are parameterized with subpopulation-specific rate data, interpolated from cumulative-germination-response curves. The purpose of this study was to evaluate the relative accuracy of three-dimensional models for predicting cumulative germination response to temperature. Three-dimensional models are relatively more efficient to implement than two-dimensional models and can be parameterized directly with measured data. METHODS: Seeds of four rangeland grass species were germinated over the constant-temperature range of 3 to 38 degrees C and monitored for subpopulation variability in germination-rate response. Models for estimating subpopulation germination rate were generated as a function of temperature using three-dimensional regression, statistical gridding and iterative-probit optimization using both measured and interpolated-subpopulation data as model inputs. KEY RESULTS: Statistical gridding is more accurate than three-dimensional regression and iterative-probit optimization for modelling germination rate and germination time as a function of temperature and subpopulation. Optimization of the iterative-probit model lowers base-temperature estimates, relative to two-dimensional cardinal-temperature models, and results in an inability to resolve optimal-temperature coefficients as a function of subpopulation. Residual model error for the three-dimensional model was extremely high when parameterized with measured-subpopulation data. Use of measured data for model evaluation provided a more realistic estimate of predictive error than did evaluation of the larger set of interpolated-subpopulation data. CONCLUSIONS: Statistical-gridding techniques may provide a relatively efficient method for estimating germination response in situations where the primary objective is to estimate germination time. This methodology allows for direct use of germination data for model parameterization and automates the significant computational requirements of a two-dimensional piece-wise-linear model, previously shown to produce the most accurate estimates of germination time.     

ESS germination strategies in randomly varying environments. I. Logistic-type models

ESS germination strategies are studied in a model of annual plant population dynamics in a randomly varying environment. The possible strategies are different values of the annual germination fraction G, either constant over time or varying in response to a "cue" correlated with upcoming environmental conditions. The model generalizes D. Cohen's model (1966, J. Theor. Biol. 12, 119-129; 1968, J. Ecol. 56, 219-228) by allowing density-dependent per capita seed yields. ESSs are characterized in terms of the resulting harmonic mean growth rate of population density. The ESS criterion cannot be solved analytically, but qualitative relationships between the value of the ESS and other population parameters are obtained, and environments in which 100% germination is an ESS are identified. Some explicit predictions of the theory are summarized and compared with ideas of M. Westoby (1981, Amer. Nat. 118, 882-885). The results of this study are compared with those of Cohen (op. cit.) in a companion paper.     

埋藏对65种半干旱草地植物种子萌发特性的影响

比较了半干旱草原科尔沁沙地65种植物新采种子和埋藏1年种子的萌发特性.结果表明：埋藏1年后,有38种植物的种子仍具有活力.新采种子萌发率>80%的植物种子埋藏后萌发率下降；而萌发率<80%的植物种子埋藏后萌发率均有较大幅度的提高.埋藏对种子萌发特性的影响因植物生态类群(杂草、草原植物、草甸植物、沙生植物)而异.种子埋藏1年后,多年生植物萌发率没有明显提高,而1年生植物萌发率显著提高(P<0.05).对于新采种子和埋藏1年种子萌发率间相差10%以下的物种,可根据萌发格局将植物种分为3类：埋藏促进型(如差巴嘎蒿)、埋藏抑制型(如大籽蒿)和埋藏不变型(如鹤虱).     

Predicting patterns of post-fire germination in 35 eastern Australian Fabaceae

Germination in 35 species from 15 legume genera of southeastern Australia was promoted by a heat treatment which broke the seed coatcaused dormancy. Once the critical temperature was reached, most seeds had their dormancy broken, independent of the duration of heating. Species fell into three classes according to whether their dormancy was broken by a temperature of 40, 60 or 80°C. Highest germination in all species was achieved by heating in the temperature range 80–100°C, although long durations (120 min) at 100°C caused seed death in several species. At 120°C, seeds of most species were killed at all but one minute's duration. A proportion of seeds from 7 species (Acacia myrtifolia, Pultenaea daphnoides, P. incurvata, P. linophylla, P. polifolia, Dillwynia floribunda and Sphaerolobium vimineurn) was not killed at 120°C and had their dormancy broken. This proportion varied markedly and resultant germination levels were significantly less than those at 80 and 100°C, except in S. vimineum. Between-site variations in the 4 species tested (A. myrtifolia, A. suaveolens, A. terminalis and A. ulicifolia) were small. These variations concerned: (i) the minimum temperature required to break seed dormancy in 2 species: 60°C in one population of A. myrtifolia and A. suaveolens, and 80°C in the other; and (ii) the intensity of the germination response. Duration of heating was less important than temperature as a determinant of germination. Ordination techniques revealed that results from one duration across temperatures were comparable with data from multiple durations. This has significant applications in studying rare species, where seed may be in short supply. Predicted germination levels after a moderate intensity fire should far exceed those after a low intensity fire. Little germination was predicted for many species after a low intensity fire and for one species, A. elongata, no germination was predicted. The potential role of indicator species in relation to the maintenance of species in a community is suggested.     

Correlates of inter-specific variation in germination response to water stress in a semi-arid savannah

Within arid plant communities species vary considerably in the ability to germinate under water stress. Attempts to correlate this variation with environmental gradients have remained largely inconclusive. Germinating only at high water potentials can be seen as a form of predictive germination. Predictive germination provides a fitness variance reducing mechanism and is therefore expected to show negative correlations with other variance reducing life-history attributes such as large seed size or dormancy. We predicted that differences in life-history attributes rather than edaphic gradients could explain the variation in germination responses to water stress found in arid plant communities. To test our hypothesis we determined the germination response of 28 species from the arid Kalahari savannah to a gradient of osmotic stress, expressed as the water potential needed to reduce germination by 50%. In addition, we determined the life-history variables seed mass and germination fraction and the habitat variables soil texture preference and association with acacias. The data were analysed using phylogenetically independent contrasts in a multiple regression model.Contrary to our hypothesis we found no increase in the capacity to germinate under osmotic stress with increasing seed mass and an increase with increasing germination fraction. However, we also found no significant effect of the habitat variables. This result may be explained by variation in seedling drought tolerance. Drought tolerance will also have a variance-reducing effect and can be expected to trade-off with fractional germination. Our results suggest that in arid plant communities most variation in the capacity to germinate under water stress expresses different ways to make a living under similar conditions rather than adaptations to environmental gradients.     

Provisioning of bioavailable carbon between the wet and dry phases in a semi-arid floodplain

Ecosystem functioning on arid and semi-arid floodplains may be described by two alternate traditional paradigms. The pulse-reserve model suggests that rainfall is the main driver of plant growth and subsequent carbon and energy reserve formation in the soil of arid and semi-arid regions. The flood pulse concept suggests that periodic flooding facilitates the two-way transfer of materials between a river and its adjacent floodplain, but focuses mainly on the period when the floodplain is inundated. We compared the effects of both rainfall and flooding on soil moisture and carbon in a semi-arid floodplain to determine the relative importance of each for soil moisture recharge and the generation of a bioavailable organic carbon reserve that can potentially be utilised during the dry phase. Flooding, not rainfall, made a substantial contribution to moisture in the soil profile. Furthermore, the growth of aquatic macrophytes during the wet phase produced at least an order of magnitude more organic material than rainfall-induced pulse-reserve responses during the dry phase, and remained as recognizable soil carbon for years following flood recession. These observations have led us to extend existing paradigms to encompass the reciprocal provisioning of carbon between the wet and dry phases on the floodplain, whereby, in addition to carbon fixed during the dry phase being important for driving biogeochemical transformations upon return of the next wet phase, aquatic macrophyte carbon fixed during the wet phase is recognized as an important source of energy for the dry phase. Reciprocal provisioning presents a conceptual framework on which to formulate questions about the resistance and ecosystem resilience of arid and semi-arid floodplains in the face of threats like climate change and alterations to flood regimes.     

Predicting the subject-specific primary stability of cementless implants during pre-operative planning: preliminary validation of subject-specific finite-element models

Pre-operative planning help the surgeon in taking the proper clinical decision. The ultimate goal of this work is to develop numerical models that allow the surgeon to estimate the primary stability during the pre-operative planning session. The present study was aimed to validate finite-element (FE) models accounting for patient and prosthetic size and position as planned by the surgeon. For this purpose, the FE model of a cadaveric femur was generated starting from the CT scan and the anatomical position of a cementless stem derived by a skilled surgeon using a pre-operative CT-based planning simulation software. In-vitro experimental measurements were used as benchmark problem to validate the bone-implant relative micromotions predicted by the patient-specific FE model. A maximum torque in internal rotation of 11.4 Nm was applied to the proximal part of the hip stem. The error on the maximum predicted micromotion was 12% of the peak micromotion measured experimentally. The average error over the entire range of applied torques was only 7% of peak measurement. Hence, the present study confirms that it is possible to accurately predict the level of primary stability achieved for cementless stems using numerical models that account for patient specificity and surgical variability.     

人工长白落叶松立木叶面积预估模型

叶面积影响着树木干物质的生产,进而影响树木乃至整个林分的生长,而叶面积准确估计对分析树木和林分生长具有重要作用.本研究基于黑龙江省长白落叶松人工林中76株解析木数据,分别建立枝条层面和单木层面的叶面积预估模型.结果表明: 考虑样木层次随机效应的最优枝条叶面积混合效应模型包含lnBD(BD为枝条基径)、lnRDINC(RDINC为相对着枝深度)和lnCR(CR为冠长率)3个随机效应参数,具体形式为:lnBLA=β₁+(β₂+b₂)lnBD+(β₃+b₃)lnRDINC+β₄lnDBH+β₅lnHT/DBH+(β₆+b₆)lnCR,其中:β_i和b_i分别是模型的固定效应参数和随机效应参数;DBH为树木胸高处直径;HT/DBH为树高与胸径的比值.模型的修正决定系数(R_a²)为0.90,均方根误差(RMSE)为0.5477,平均偏差(ME)为-0.03,平均绝对偏差(MAE)为0.24,预测精度(P)为91%,枝条叶面积预估模型的预估效果较好.以枝条叶面积预估模型为基础,计算树冠叶面积并建立树冠叶面积预估模型,最终形式为:lnCLA=γ₀+γ₁lnDBH+γ₂CR,其中,γ_i为模型参数.似然比检验结果(P>0.05)说明该模型不用考虑样地层次的随机效应.本研究所建立的立木树冠叶面积预估模型的决定系数(R²)为0.87,RMSE为0.3847,拟合效果好,可以很好地预测人工长白落叶松立木树冠叶面积,为以后叶面积分布和光合作用的研究提供了理论基础.     

Erythroblastosis models. II. Materno-fetal incompatibility in chimpanzee

Erythroblastosis fetalis represents a significant hazard for successful management of pregnancy in man and in marmosets, but not in crab-eating macaques. Materno-fetal blood group incompatibility in chimpanzee is described as a contributing factor in the death of an infant. The findings indicate that parental blood groups should be taken into consideration when breeding chimpanzees.     

Dormancy and impotency of cocklebur seeds II. Phase sequence in germination process

Interactions of C₂H₄, CO₂, O₂ and high temperature in stimulatinggermination of cocklebur (Xanthium pennsylvanicum Wallr.) seedswere studied and the phase sensitive to each factor during germinationwas determined. The combination of CO₂ and enriched O₂, andparticularly that of C₂H₄ and enriched O₂, much more effectivelystimulated germination than CO₂ and C₂H₄. At low temperature,however, the cooperation of CO₂ and enriched O₂ was lost andonly the effect of C₂H₄ in combination with CO₂ or enrichedO₂ remained. Carbon dioxide stimulated C₂H₄ production and induced germinationwhen it was applied in the first period of water imbibition,corresponding to the passive thrust forming phase. C₂H₄ becameeffective after the CO₂-responsive phase. In contrast, bothO₂ enrichment and high temperature became increasingly effectivewith the imbibition times. Anaerobiosis applied during the firstperiod of the germination process showed no inhibitory effect,whereas CO₂ and C₂H₄ were stimulative even under such a condition. (Received August 26, 1974; )