Quantal Response of Fruit and Seed Germination Rate in Quercus robur L. and Castanea sativa Mill, to Constant Temperatures and Photon Dose

A linear relationship between constant temperatures in the sub-optimaltemperature range and germination rate is shown in both Quercusrobur L and Castanea sativa Mill germinated under nominal darkconditions. The mean base temperature was interpolated for Qrobur as 0 8 ? or 2-4 ?, depending on seed lot provenance, andfor C. sativa as 1 -4? The optimum temperature for germinationin Q. robur was about 20? compared with around 28 ? in C. sativaOver the sub-optimal temperature range the distribution of thermaltimes was log-normal for each population studied their spreadvarying both between Q robur seed lots and between species However,in C. sativa germinated close to the mean base temperature,the distribution in thermal times was reduced Thermal timesto germination were decreased in Q. robur and C sativa by approximately0 3 and 0-5 log-units, respectively, when the pericarp was removed,i.e in the seeds, but the sensitivity of the response remainedrelatively unaltered In both species the germination rate was the same when nominaldark or safe green light conditions were employed during thegermination test. However, at 21 ? Q robur exhibited the highirradiance reaction (HIR) at photon doses above 30mmol m^–2d^–1. HIR first affected the germination rate by an inhibitionof radicle extension The sensitivity of the response to thermaltime was reduced as photon dose increased. This photo-inhibitionwas exacerbated at supra-optimal temperatures. In contrast,C. sativa germination rate at 26 ? was little influenced bylight at a photon dose of 752 mmol m^–2 d^–1 Key words: Seed germination rate, temperature, thermal time, light, photon dose     

Quantal Response of Seed Germination in Seven Genera of Cruciferae to White Light of Varying Photon Flux Density and Photoperiod

The response of the germination of seeds of Barbarea vema (Mill.)Aschers, Brassica chinensis L., Brassica juncea (L.) Czern.& Coss., Brassica oleracea L. var. gongylodes L., Camelinasaliva (L.) Crantz, Eruca saliva Mill., Lepidium sativum L.,Nasturtium officinale R. Br., and Rorippa palustris (L.) Besserto white fluorescent light of different photon flux densitiesapplied for different daily durations in a diurnal alternatingtemperature regime of 20 °C/30 °C (16 h/8 h) was quantifiedby linear relations between probit percentage germination andthe logarithm of photon dose, the product of photon flux densityand duration. The low energy reaction, in which increasing dosepromotes germination, was detected in all the seed populationsbut in Barbarea vema and Brassica Juncea the lowest photon doseapplied (10^–5–2 and 10^{–5 7} mol m^–2 d^–1,respectively) was sufficient to saturate the response. Comparisons,where possible, between photoperiods demonstrated reciprocity,i.e. germination was proportional to photon dose irrespectiveof photoperiod, for the low energy reaction in Brassica oleracea(1 min d^–1 to 1 h d^–1), Camelina saliva (1 min d^–1to 8 h d^–1), Eruca saliva (1 min d^–1 to 24 h d^–1),Lepidium sativum (I min d^–1 to 8 h d^–1) and Rorippapalustris (1 min d^–1 to 8 h d^–1), but not in Brassicachinensis and Nasturtium officinale. The high irradiance reaction,in which increasing dose inhibits germination, was detectedin Barbarea vema, Brassica chinensis, Brassica juncea, Brassicaoleracea, and Camelina saliva. The minimum dose at which inhibitionwas detected was lO^–0–3 mol m^–2 d^–1.These results are discussed in the context of devising optimallight regimes for laboratory tests intended to maximize germination The response of germination to photon dose was also quantifiedwith 3 x 10^–4 M GA₂, co-applied (Brassica chinensis, Camelinasaliva, and Lepidium sativum) and with 2 x 10^–2 M potassiumnitrate co-applied (Brassica chinensis). In the latter casepotassium nitrate had no effect in the dark and inhibited germinationin the light, but GA₂, promoted germination substantially inall three species. Variation amongst seeds in the minimum photondose required to stimulate germination was not affected by co-applicationof GA₂, in Brassica chinensis and Camelina saliva, whereas seedsof Lepidium salivum showed a narrower distribution of sensitivitiesto the low energy reaction in the presence of GA₂ Barbarea vema (Mill.) Aschers, Brassica chinensis L., Brassica juncea (L.) Czern. & Coss., Brassica oleracea L. var. gongylodes L., Camelina saliva (L.) Crantz, Eruca saliva Mill., Lepidium satiaum L., Nasturtium officinale R. Br., Rorippa palustris (L.) Besser, Cruciferae, light, gibberellic acid, seed germination, seed dormancy     

Response of Seed Germination in Three Genera of Compositae to White Light of Varying Photon Flux Density and Photoperiod

Various photon doses (net number of photons per unit area perday), provided by varying both photon flux density and photoperiod,were applied to imbibing seeds of seven lots of four speciesof Compositae in various germination test regimes. In all fourspecies germination was dependent upon photon dose, the productof photon flux density and daily duration of exposure. The responsewas quantified by linear relations between the probit of percentagegermination and the logarithm of photon dose. In general, photonflux density and photoperiod only influenced the stimulationof germination by the low energy reaction indirectly (as factorsof daily photon dose), whereas there was a tendency for photoperiodto have a direct influence on the inhibition of germinationby the high irradiance reaction. Reducing the germination testtemperature from 25?C to 20?C and 15?C not only increased thedark germination of L. sativa L., but also broadened the photondose range at which full germination occurred by reducing theminimum value necessary for the germination of the most dormantseeds, and increasing the maximum value which failed to inhibitthe germination of any seeds. Differences between L. sativaand L. serriola L. in the response of germination to white lightwere only quantitative, rather than qualitative. The singlemost promotory dose for all four species was 3 ? 10^–3mol m^–2 d^–1, although the inhibitory action of dosesup to 10– mol m– d– was generally only slight. Key words: Light, seed germination, seed dormancy, Compositae     

The effect of fermentation on the growth and survival ofSalmonella typhimurium,Staphylococcus aureus,Bacillus cereus andPseudomonas aeruginosa in fermenting tef (Eragrostis tef)

Summary Growth ofSalmonella typhimurium, Staphylococcus aureus, Bacillus cereus andPseudomonas aeruginosa was inhibited when the pH of fermenting tef approached 5.0, 5.0, 5.5 and 5.0, respectively. However the test organisms grew in far more acidic conditions in broth than in fermenting tef and this is due to antimicrobial substance(s) being produced by some of the lactic acid bacteria. Except forBacillus cereus spores, all the test organisms were heat-inactivated during the baking process of the final tef injera.

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Effet de la fermentation sur la croissance et la survie de Salmonella typhimurium, Staphylococcus aureus, Bacillus cereus et Pseudomonas aeruginosa dans le tef (Eragrostis tef) en fermentation

Résumé La croissance deSalmonella typhimurium, deStaphylococcus aureus, deBacillus cereus et dePseudomonas aeruginosa est inhibée lorsque les pH du tef en fermentation approchent respectivement 5:0, 5.0, 5.5 et 5.0. Toutefois, les organismes tests croissent dans des conditiones bien plus acides que dans le tef en fermentation. Ceci est dû à des substances antimicrobiennes produites par certaines bactéries lactiques. A l'exception des spores deBacillus cereus, tous les organismes tests sont inactivés par la chaleur durant le processus de cuisson.

     

In Silico Approach for Unraveling the Structural and Functional Roles of NF-X1-Like Proteins in Underutilized Cereal Eragrostis tef

Biology Bulletin - Eragrostis tef is a increasingly becoming popular in different parts of the world as cereal due to its wide acceptability in extreme environmental conditions where most other...     

The Interactive Effects of Phytochrome, Nitrate and Thiourea on the Germination Response to Alternating Temperatures in Seeds of Ranunculus sceleratus L.: A Quantal Approach

Probert, R. J., Gajjar, K. H. and Haslam, I. K. 1987. The interactiveeffects of phytochrome, nitrate and thiourea on the germinationresponse to alternating temperatures in seeds of Ranunculussceleratus L.: A quantal approach.—J. exp. Bot. 38: 1012–1025. The interactive effects of phytochrome, potassium nitrate andthiourea on the germination response to alternating temperaturesin achenes (seeds) of Ranunculus sceleratus L. were studied.Using thermogradient bars, high levels of germination were recordedover a broad range of alternating temperatures providing seedsreceived daily irradiations. Reduced germination in temperaturecycles with a relatively long warm phase was related to thelevel of the active form of phytochrome (Pfr). Dose-responseexperiments to red light (R) and temperature shifts showed thatthe actions of Pfr and alternating temperatures were interdependent.Maximum germination was recorded when intermittent pulses ofR were combined with daily 4 h temperature shifts from 16°Cto 26°C. Whilst probit analysis showed that potassium nitrateand thiourea both increased population sensitivity to temperatureshifts, thiourea was a more potent stimulant. Although the effectof both chemicals was dependent on phytochrome photo-equilibriumthe threshold level of Pfr required for thiourea action wasclearly much lower than that required for nitrate action. Thioureapotentiated a response to daily temperature shifts even whenPfr was at a low, normally inhibitory level. These results indicatedifferent mechanisms of action for potassium nitrate and thioureain relation to phytochrome controlled seed germination. Key words: Phytochrome, nitrate, thiourea, alternating temperatures, germination     

Development of ABA Antagonists to Overcome ABA- and Low Temperature-Induced Inhibition of Seed Germination in Canola,Lentil, and Soybean

ABA antagonists have potential application as growth regulators to improve germination and seedling growth at low temperatures for oilseeds and pulses grown in regions with short seasons such as those in western Canada. Towards development of practical ABA antagonists, a series of 3′-substituted ABA analogs were synthesized and screened in seed germination assays in canola (Brassica napus), lentil (Lens culinaris), and soybean (Glycine max) at low temperature and in overcoming exogenous ABA. The most promising analog, ABA 1009, was selected for further germination testing of dose responses in canola, lentil, and soybean. Analog ABA 1009 at 100 µM was effective in overcoming ABA (10 µM)-induced inhibition for canola, lentil, and soybean germination at ambient temperature, and also promoted germination at low temperature for canola (5 °C).

     

Regulation of Electron Transport in Photosystems I and II in C3, C3-C4, and C4 Species of Panicum in Response to Changing Irradiance and O2 Levels

Regulation of the quantum yields of linear electron transport and photosystem II photochemistry ([phi]II) with changing irradiance and gas-phase O2 concentration was studied in leaf tissue from Panicum bisulcatum (C3), Panicum milioides (C3-C4), and Panicum antidotale (C4) at 200 [mu]bars of CO2 and 25[deg]C using infrared gas analysis and chlorophyll fluorescence yield measurements. When the O2 level was increased from 14 to 213 mbars at high irradiance, [phi]II increased by as much as 115% in P. bisulcatum but by no more than 17% in P. antidotale. Under the same conditions [phi]II increased to an intermediate degree in P. milioides. Measurements of accumulation of the photooxidized form of the photosystem I reaction center (P700+) based on the light-dependent in vivo absorbance change at 830 nm indicate that the steady-state concentration of P700+ varied in an antiparallel manner with [phi]II when either the irradiance or O2 concentration was changed. Hence, O2-dependent changes in [phi]II were indicative of variations in linear photosynthetic electron transport. These experiments revealed, however, that a significant capacity was retained for in vivo regulation of the apparent quantum yield of photosystem I ([phi]I) independently of [phi]II+ Coordinate regulation of quantum yields of photosystems I and II (expressed as [phi]I:[phi]II in response to changing irradiance and O2 level differed markedly for the C3 and C4 species, and the response for the C3-C4 species most closely resembled that observed for the C4 species. The fraction of total linear electron transport supporting photorespiration at 213 mbars of O2 was negligible in the C4 species and was 13% lower in the C3-C4 species relative to the C3 species as calculated from fluorescence and gas-exchange determinations. At high photon-flux rates and high O2 concentration, the potential benefit to light use for net CO2 uptake arising from lower photorespiration in P. milioides was offset by a reduced capacity for total CO2- and O2-dependent noncyclic electron transport in this species compared with P. bisulcatum.     

Significance of the Leaf Area Ratio in Hevea brasiliensis Under High Irradiance and Low Temperature Stress

Adjustment in leaf area : mass ratio called leaf area ratio (LAR) is one of the strategies to optimize photon harvesting. LAR was recorded for 10 genotypes of Hevea brasiliensis under high irradiance and low temperature and the genotypes were categorized into two groups, i.e. high LAR and low LAR types. Simultaneously, the growth during summer as well as winter periods, photosynthetic characteristics, and in-vitro oxidative damage were studied. Low LAR (19.86±0.52 m² kg^–1) types, recorded an average of 18.0 % chlorophyll (Chl) degradation under high irradiance and 7.1 % Chl degradation under low temperature. These genotypes maintained significantly higher net photosynthetic rate (P _N) of 10.4 mol(CO₂) m^–2 s^–1 during winter season. On the contrary, the high LAR (24.33±0.27 m² kg^–1) types recorded significantly lower P _N of 4 mol(CO₂) m^–2 s^–1 and greater Chl degradation of 37.7 and 13.9 % under high irradiance and low temperature stress, respectively. Thus LAR may be one of the physiological traits, which are possibly involved in plant acclimation process under both stresses studied.     

胡杨种子萌发对温光条件和盐旱胁迫的响应特征

以胡杨(Populus euphratica)种子为材料,分别设置光照(连续光照、12h光照/12h黑暗、连续黑暗)温度(10/15℃、15/20℃、20/25℃、25/30℃、30/35℃和35/40℃)试验、PEG6000渗透胁迫(0、-0.1、-0.2、-0.4、-0.6、-0.8、-1.0、-1.2、-1.4和-1.6MPa)试验和NaCl胁迫(0、0.05、0.10、0.20、0.30、0.40、0.60和0.80mol/L)试验,考察室内种子萌发对温度、光照和盐旱胁迫的敏感性,揭示胡杨种子萌发阶段对生境资源的适应策略。结果显示:(1)胡杨种子在温度(10℃/15℃~35℃/40℃)与连续光照、连续黑暗和12h光照/12h黑暗组合处理条件下均能萌发,且最终种子萌发率均能达到77%以上;3种光照条件下,种子萌发的最适温度范围均为25℃/30℃~30℃/35℃,在该温度范围种子萌发表现出快速、集中的特点,且3种光照条件对种子萌发的影响无显著差异。(2)胡杨种子可以在-1.4~0 MPa渗透势溶液中萌发,而在-1.0~0 MPa间最终萌发率均达到90%以上,且相互之间无显著差异;但当渗透势低于-0.4 MPa时胡杨种子萌发进程和萌发速率受到显著影响,当溶液渗透势低于-1.2 MPa时种子萌发受到显著抑制。(3)胡杨种子可以在0~0.80mol/L NaCl溶液中萌发,而最终萌发率、萌发速率均随着NaCl溶液浓度的增高逐渐降低,但在0~0.20mol/L范围内无显著差异;当NaCl溶液浓度大于0.20mol/L时,种子最终萌发率、种子萌发进程和萌发速率均受到显著抑制,萌发高峰期逐渐向后推移。研究结果表明,胡杨种子萌发时温度比较宽泛,对光照无严格要求,适宜温度下萌发快速集中,且萌发时对盐旱胁迫具有一定程度的忍耐性。这些特性有助于胡杨种子充分利用有限的水分条件而快速完成萌发,是胡杨种子萌发对干旱荒漠地区干旱少雨环境的一种生态适应策略。     

A Guide to Bivariate Ideas of Quantal Response Analysis,as Applied in Biometrics and Econometrics

A review is provided of the several bivariate generalisations of quantal response analysis that have appeared in the biometric and econometric literatures. There are three main types: (i) where a binary outcome is the result of two stimulants, and thus the bivariate distribution of the thresholds for response is relevant; (ii) where three or more alternative outcomes may arise from a single stimulant; and (iii) where the response itself is bivariate (i.e., two types of response may simultaneously be observed).     

Response to Salinity in Legume Species: An Insight on the Effects of Salt Stress during Seed Germination and Seedling Growth

The process of soil salinization and the preponderance of saline water sources all over the world represent one of the most harmful abiotic stress to plant growth. This pointed to the importance of obtaining plants which are tolerant or resistant to salt, considering that projection of climate change for the coming years indicate an increase in temperature and rain scarcity. In the current study, the effect of NaCl was investigated on germinating seeds of Lathyrus sativus L., Vicia sativa L., Vigna radiata L. R.Wilczek and Vigna unguiculata L. Walp., by combining physiological, biochemical, biostatistical and ultrastructural analyses. Our results revealed that germination was not influenced by high NaCl concentrations, while seedling growth was affected even at low NaCl concentrations, probably due to an alteration in water uptake and in organic matter biosynthesis. Nevertheless, the synthesis of antioxidant enzymes, phenolic acids and flavonoids was registered in all species, which tended to cope with the increasing salt stress, allowing a response mechanism such as cytoplasm detoxification and cellular turgor maintenance. Besides, the ultrastructural analysis evidenced plasmolyzed cells close to cells with a normal ultrastructure with no appreciable differences among the species. This research deeply investigates the mechanism of salt-stress response focusing on species never tested before for their possible tolerance to salinity.     

赤霉素、钙和甜菜碱对小桐子种子萌发及幼苗抗低温和干旱的影响

小桐子(Jatropha curcas L.)属大戟科( Euphorbiaceae)麻疯树属(Jatropha L.)的能源植物。分别以小桐子种子和幼苗为实验材料,研究了不同浓度的赤霉素和CaCl₂单独处理,以及不同浓度的赤霉素、CaCl₂和甜菜碱组合处理对小桐子种子萌发及幼苗抗低温和干旱的影响。结果表明:分别用10 mg/L赤霉素和10 mmol/L CaCl₂处理小桐子种子,不仅可以提高其在正常萌发条件(26℃)、低温(18℃)和干旱胁迫(5% PEG6000)下的发芽率,还可缓解小桐子幼苗在低温(2℃)或干旱胁迫(25% PEG6000)下电解质渗漏率的增加和丙二醛(MDA)的积累。用10 mg/L赤霉素、5 mmol/L CaCl₂和15 mmol/L甜菜碱组合处理小桐子种子,也可进一步增强其在正常萌发(26℃)、低温(18℃)和干旱胁迫(5% PEG6000)条件下种子的发芽率,以及缓解小桐子幼苗在低温(2℃)或干旱胁迫(25% PEG6000)下电解质渗漏率的增加和MDA的积累,表明10 mg/L赤霉素和10 mmol/L CaCl₂分别处理或10 mg/L赤霉素、5 mmol/L CaCl₂和10 mmol/L甜菜碱组合处理可提高小桐子种子在低温和干旱胁迫下的发芽率,以及提高小桐子幼苗对低温和干旱胁迫的抵抗能力。     

Xanthophyll Cycle and Inactivation of Photosystem Ⅱ Reaction Centers Alleviating Reducing Pressure to Photosystem Ⅰ in Morning Glory Leaves under Short-term High Irradiance

Under 30-min high irradiance （1500μmol m^-2 s^-1）, the roles of the xanthophyll cycle and D1 protein turnover were investigated through chlorophyll fluorescence parameters in morning glory （Ipomoea setosa） leaves, which were dipped into water, dithiothreitol （DTT） and lincomycin （LM）, respectively. During the stress, both the xanthophyll cycle and D1 protein turnover could protect PSI from photoinhibition. In DTT leaves, non-photochemical quenching （NPQ） was inhibited greatly and the oxidation level of P700 （P700^＋） was the lowest one. However, the maximal photochemical efficiency of PSII （Fv/Fm） in DTT leaves was higher than that of LM leaves and was lower than that of control leaves. These results suggested that PSI was more sensitive to the loss of the xanthophyll cycle than PSII under high irradiance. In LM leaves, NPQ was partly inhibited, Fv/Fm was the lowest one among three treatments under high irradiance and P700^＋ was at a similar level as that of control leaves. These results implied that inactivation of PSII reaction centers could protect PSI from further photoinhibition. Additionally, the lowest of the number of active reaction centers to one inactive reaction center for a PSII cross-section （RC/CSo）, maximal trapping rate in a PSll cross-section （TRo/CSo）, electron transport in a PSll cross-section （ETo/CSo） and the highest of 1-qP in LM leaves further indicated that severe photoinhibition of PSII in LM leaves was mainly induced by inactivation of PSII reaction centers, which limited electrons transporting to PSh However, relative to the LM leaves the higher level of RC/CSo, TRo/CSo, Fv/Fm and the lower level of 1-qP in DTT leaves indicated that PSI photoinhibition was mainly induced by the electron accumulation at the PSI acceptor side, which induced the decrease of P700^＋ under high irradiance.     

Variation in response of accessions of minor millets,Pennisetum americanum (L.) Leeke (pearl millet) and Eleusine coracana (L.) Gaertn (finger millet), and Eragrostis tef (Zucc.) Trotter (tef) to salinity in early seedling growth

The response to increasing NaCl concentration of seedlings of 25 accessions of Ethiopian land races of each of Pennisetum americanum (L.) Leeke (pearl millet) and Eleusine coracana (L.) Gaertn (finger millet), and 15 accessions of Eragrostis tef (Zucc.) Trotter (tef), was examined after two week's growth in NaCl solution culture. Although increasing NaCl concentration significantly reduced seedling root lengths, there was considerable variation within, and between accessions within each species.Analysis based upon a non-linear least square inversion method, using root length data, revealed significant differences in accessions of P. americanum and E. tef on the basis of the estimated salinity threshold, C _t , the NaCl concentrations at which root length begins to decrease. C _t did not differ significantly between E. coracana accessions. Estimates of C₅₀ and C₀, mininum concentrations causing a 50% decrease in root length, and zero root growth respectively, revealed differences between and within accessions for all three species. Overall, finger millet was more tolerant than tef, which was more tolerant than pearl millet. There is clear evidence that differences in tolerance are genetically based from broad sense heritability estimates.     

Responses of the Antioxidant Systems and Xanthophyll Cycle in Phaseolus vulgaris to the Combined Stress of High Irradiance and High Temperature

Changes in the activities of enzymes involved in scavenging active oxygen species were followed after exposing bean seedling leaves (Phaseolus vulgaris L.) to various cross stresses of irradiance and temperature. The activities of superoxide dismutase (SOD, EC 1.15.1.1) and ascorbate peroxidase (AsAPOD, EC 1.11.1.11) increased to different extent with prolonged irradiation of the leaves, and were stimulated by high temperature (HT). The activity of catalase (CAT, 1.11.1.6) decreased when exposed to strong irradiance (HI), and the decrease was further exacerbated when HI was combined with HT. CAT activity was more sensitive to HT than to HI. Ascorbate (AsA) content slightly decreased and then increased during the treatment of HI, but decreased under the cross stress of HI and HT. On the contrary, glutathione (GSH) content increased all the time during various treatments of irradiance and temperature. The increase in the combined stress was even more pronounced. Irradiance is the major reason in triggering the operation of xanthophyll cycle, which was difficult to be started by HT. The antioxidant systems tended to be inactivated with prolonged exposure to the cross stress of HI and HT. The de-expoxidated state of xanthophyll cycle, however, was increasing all the time, which indicated that the zeaxanthin-dependent thermal dissipation was one major energy dissipation pathway during the cross stress of HI and HT.     

Phytochrome A Mediates the Promotion of Seed Germination by Very Low Fluences of Light and Canopy Shade Light in Arabidopsis

Seeds of the wild type (WT) and of the phyA and phyB mutants of Arabidopsis thaliana were exposed to single red light (R)/far-red light (FR) pulses predicted to establish a series of calculated phytochrome photoequilibria (Pfr/P). WT and phyB seeds showed biphasic responses to Pfr/P. The first phase, i.e. the very-low-fluence response (VLFR), occurred below Pfr/P = 10-1%. The second phase, i.e. the low-fluence response, occurred above Pfr/P = 3%. The VLFR was similarly induced by either a FR pulse saturating photoconversion or a subsaturating R pulse predicted to establish the same Pfr/P. The VLFR was absent in phyA seeds, which showed a strong low-fluence response. In the field, even brief exposures to the very low fluences of canopy shade light (R/FR ratio < 0.05) promoted germination above dark controls in WT and phyB seeds but not in the phyA mutant. Seeds of the phyA mutant germinated normally under canopies providing higher R/FR ratios or under deep canopy shade light supplemented with R from light-emitting diodes. We propose that phytochrome A mediates VLFR of A. thaliana seeds.     

Changes in the Activity of Some Hydrolases, Peroxidase, and Catalase in the Rice Seed during Germination

A study was made of the changes in activity of enzymes involved in the breakdown of stored phytin, lipid, and hemicellulose in the aleurone layer of rice seed (Oryza sativa L., variety IR8) during the 1st week of germination in the light. Enzyme assays were made on crude extracts from degermed seed, and activities were expressed on a per seed basis. Phytase activity increased within the 1st day of germination. The increase in activity of most other enzymes—phosphomonoesterase, phosphodiesterase, esterase, lipase, peroxidase, catalase, β-glucosidase, and α- and β-galactosidase—closely followed the increase in protein content. Their peak activities occurred by the 5th to the 7th day. Some enzymes, such as β-1, 3-glucanase and α-amylase, continued to increase in activity after the 7th day. Phytase, β-1, 3-glucanase, and α-amylase followed a similar sequence of production in embryoless seed halves incubated in 0.12 μM gibberellin A₃, but the production of lipase was delayed.