Papaver somniferum

Propagation

Germination

There is some disagreement on the ideal germination temperature for somniferum. Part of this is due to differences in varietal responses and part is due to nonstandardized test conditions.[3]

The ideal germination temperature varies slightly by cultivar but is generally even in the range of 15-20°C. Germination at 30-35°C is massively suppressed in most cultivars.[1][4][3] The variation seems to be an adaptation to region specific growing cycles: winter or spring poppies. Higher heat tolerance is typical of the varieties adapted to germinate in the fall and overwinter.[3]

There is a slight advantage to placing seeds between two sheets of filter paper (“blotters”) compared to placement on top of a single sheet but only at 4°C. The author speculates that this effect might be due to evaporative cooling in “icebox” conditions since it was not present in any other test group. [1]

There is a clear trend of increasing rapidity of germination with increasing temperature at the expense of a lower final germination rate at the high end of temperature.[1][4]

Germination at an alternating temperature of 20/30°C (mean temperature 22.5°C) is equivalent to a constant temperature of 22.5°C.[1]

Harrington recommends germination on top of filter paper of somniferum at 15°C. The test is complete by eight days after sowing.[1]

Unlike P. bracteatum[2], P. somniferum does not require light to germinate.[4][2] Further, illumination with red and far-red lights had no effect on germination of somniferum.[2]

Somniferum seeds seem to be sensitive to pH. Germination at pH 5 and pH 9 were significantly reduced compared to pH 7 (82%, 64%, and 97%, respectively).[4]

Somniferum germination is restricted in water submerged conditions. The effect was partially ameliorated by the inclusion of growing media.[4] This suggests that soaking seeds in water in germination conditions (e.g. refrigerator) will result in lower germination.

[3] [5]

[6] [7] [8] [9] [10] [11]

[12] [13] [14] [15] [16] [17] [18] [19]

Vegetative

In-Vitro

Cultivation

Harvest

130-170 days to harvest [21]

Yield

[22] [23] [24] [25] [26] [27] [28] [20]

Soilless

Soil

Seedlings can be transplanted after 30 days when they are 8-12cm in height with about an 80% survival rate.[20]

[22] [24] [28]

[29] [20]

Fertilization

| type | rate | time | note | reference | | —- | —- | —- | —- | ——— |

Somniferum is sensitive to cadmium accumulation.[29]

[28] [29] [20] [30]

Temperature

[3]

Lighting

| fixture type | photoperiod | illumination | note | reference | | ———— | ———– | ———— | —- | ——— | | | | | | |

Pests

Ecology

Morphology

[25] [28]

Roots

Stem

Leaves

Inflorescence

Seeds

Phytochemistry

[22] [23] [24]

Infraspecific Variation

[23] [26] [32] [27] [33]

Biosynthesis

Distribution

Timecourse

[24] [20]

Improvement

[32] [27]

Identification

[5] [32] [24] [22] [33]

Inheritance

[26] [34] [35] [30]

Methods

[34] [35] [30]

History & Society

Work Log

Bibliography

1. Harrington, Geo T, Optimum Temperatures for Flower Seed Germination, Botanical Gazette, vol. 72, no. 6, pp. 337--358, December 1921. url: https://www.journals.uchicago.edu/doi/pdfplus/10.1086/332941. Abstract
   The proper conditions for the germination of flower seeds is a subject upon which but little work has been published. During the spring of I9I2, preliminary work was done in the seed laboratory of the United States Department of Agriculture on the temperature conditions best suited for the germination of a few of the more common flower seeds. During the winter and spring of I9W3-I9W4, further work was done with the same species investigated in I9I2, and with a few additional species. The publication of the results has been unavoidably delayed for several years. In the meantime, the recommendations included herein have been followed by the seed laboratory with good results.
2. Bare, C. E. and Toole, V. K. and Gentner, W. A., Temperature and Light Effects on Germination of Papaver Bracteatum Lindl., P. Orientale L., and P. Somniferum L., Planta Medica, vol. 34, no. 6, pp. 135--143, September 1978. doi: 10.1055/s-0028-1097425. Abstract
   Thieme E-Books \& E-Journals
3. Kamkar, Behnam and {Jami Al-Alahmadi}, Majid and {Mahdavi-Damghani}, Abdolmajid and Villalobos, Francisco J., Quantification of the Cardinal Temperatures and Thermal Time Requirement of Opium Poppy (Papaver Somniferum L.) Seeds to Germinate Using Non-Linear Regression Models, Industrial Crops and Products, vol. 35, no. 1, pp. 192--198, January 2012. doi: 10.1016/j.indcrop.2011.06.033. Abstract
   The response of plant development rate (including germination rate) to temperature might be described as a non-linear function. We compared 3 non-linear regression models (Dent-like, segmented and beta) to describe the germination rate-temperature relationships of opium poppy (Papaver somniferum L.) over 6 constant temperatures to find cardinal temperatures and thermal time required to reach different germination percentiles. Two replicated experiments were performed with the same temperatures. An iterative optimization method was used to calibrate the models and different statistical indices (mean absolute error, coefficient of determination (R2), intercept and slope of the regression equation of predicted vs. observed germination rate) were applied to compare their performance. The segmented was found to be the best model to predict germination rate (R2=0.92, MAE=0.0011 and CV of 1.4–3.6\%). Estimated cardinal temperatures were similar for different germination percentiles (P{$<$}0.05). Base on the model outputs, the base, the optimum and the maximum temperatures for germination were estimated as 3.02, 27.36 and 36.31°C. The thermal time required to reach 50 and 95\% germination was 57.27 and 87.55degree-days, respectively. Model predictions of the time required for seed germination agreed reasonably well with the observed times (MAE=0.56 day, R2=0.887). All model parameters may be readily used in crop simulation models.
4. Lee, Ho Joon and Kim, Sung Jong and Kim, Seon Ho and Kim, Se Young, Effects of Temperature, pH, Light and Degree of Oxygen Supply on the Germination of Papaver somniferum L. Seeds, The Korean Journal of Ecology, vol. 9, no. 1, pp. 19--24, 1986. url: https://koreascience.kr/article/JAKO198611919964895.page. Abstract
   Effects of temperature, pH, light-and-dark and degree of oxygen supply on germination of Papaver somniferum L. seeds were investigated. The optimum temperature of the germination was 2{$<$}TEX{$>\$$}0\textasciicircum\{\textbackslash circ\}C\${$<$}/TEX{$>$}, but the germination was decreased as temperatures changed to the upper and lower. The largest elongation of both radicle and hypocotyl also occurred at 2{$<$}TEX{$>\$$}0\textasciicircum\{\textbackslash circ\}C\${$<$}/TEX{$>$}. The best germination occurred at pH 7 but this decreased suddenly the above and below of that pH value. Although no effect of light-and-dark on the germination was, this was restricted under limited. oxyten supply in terms of underwater germination as compared with abovewater one.
5. Kumar, Birendra and Verma, A K and Ram, G and Singh, H P and Lal, R K, Seed Germination of Five Elite Genotypes of Papaver Somniferum, Journal of Tropical Medicinal Plants, vol. 11, no. 1, pp. 6, 2010. Abstract
   The opium poppy Papaver somniferum is of immense pharmaceutical importance. Seeds are rich source of the fatty oil, protein and also have considerable importance as culinary item in India. Details of seed germinability of five opium poppy varieties viz. Sanchita, Vivek, Sampada, Sujata and Sweta under two test conditions viz. in-situ (Petri dish) and ex-situ (soil) and the period (i.e. first and final count day) of seed germination are outlined.
6. Benková, Michaela, Comparison of Germinability Selected Seeds After Ten Years Storage Porovnanie Klíčivosti Semien Niektorých Druhov Rastlín Po 10 Rokoch Uskladnenia, 2008. url: https://www.semanticscholar.org/paper/COMPARISON-OF-GERMINABILITY-SELECTED-SEEDS-AFTER-PO-Benkov%C3%A1/556f33b1236182fd7ecd1ee37c6f001bd7c2ba8d. Abstract
   Ing. Michaela Benková, PhD., Slovak Agricultural Research Centre – Research Institute of Plant Production, 921 68 Piešťany, Bratislavská 122. E-mail:benkova@vurv.sk RNDr. Mária Žáková, CSc., Slovak Agricultural Research Centre – Research Institute of Plant Production, 921 68 Piešťany, Bratislavská 122. E-mail:zakova@vurv.sk Nine species (Phaseolus vulgaris L., Zea mays L., Lycopersicon esculentum MILL., Pisum sativum L., Papaver somniferum L., Vicia sativa L., Linum usitatissimum L., Capsicum annum L. and Beta vulgaris L.) representing 262 accessions mostly of Slovak and former Czechoslovakia origin were monitored after ten years of storage. Genetic resources were stored in the genebank (GB) in Piešťany in the base collection at –17°C in a glass container with silica gel. Viability was monitored by conducting a germination test on a fixed sample, one hundred seeds were tested in two replications. A variety of results was obtained, using the paired T-test. Some species showed a significant increase in percentage germination (Pisum sativum L., Papaver somniferum L. and Vicia sativa L.) and some showed a decrease or no change in percentage germination (Phaseolus vulgaris L., Zea mays L., Lycopersicon esculentum MILL., Linum usitatissimum L., Capsicum annum L. and Beta vulgaris L.) Within the different species, consistent results were not always obtained, with individual accessions sometimes showing varied germination results.
7. Bonilla, A. and Sarria, A. L. F. and Algar, E. and Muñoz Ledesma, F. J. and Ramos Solano, B. and Fernandes, J. B. and Gutierrez Mañero, F. J., Microbe Associated Molecular Patterns from Rhizosphere Bacteria Trigger Germination and Papaver Somniferum Metabolism under Greenhouse Conditions, Plant Physiology and Biochemistry, vol. 74, pp. 133--140, January 2014. doi: 10.1016/j.plaphy.2013.11.012. Abstract
   Ten PGPR from different backgrounds were assayed on Papaver somniferum var. Madrigal to evaluate their potential as biotic elicitors to increase alkaloid content under the rationale that some microbe associated molecular patterns (MAMPs) are able to trigger plant metabolism. First, the 10 strains and their culture media at two different concentrations were tested for their ability to trigger seed germination. Then, the best three strains were tested for their ability to increase seedling growth and alkaloid levels under greenhouse conditions. Only three strains and their culture media enhanced germination. Then, germination enhancing capacity of these best three strains, N5.18 Stenotrophomonas maltophilia, Aur9 Chryseobacterium balustinum and N21.4 Pseudomonas fluorescens was evaluated in soil. Finally, the three strains were applied on seedlings at two time points, by soil drench or by foliar spray. Photosynthesis was measured, plant height was recorded, capsules were weighted and alkaloids analyzed by HPLC. Only N5.18 delivered by foliar spray significantly increased plant height coupled to an increase in total alkaloids and a significant increase in opium poppy straw dry weight; these increases were supported by a better photosynthetic efficiency. The relative contents of morphine, thebaine, codeine and oripavine were affected by this treatment causing a significant increase in morphine coupled to a decrease in thebaine, demonstrating the effectivity of MAMPs from N5.18 in this plant species. Considering the increase in capsule biomass and alkaloids together with the acceleration of germination, strain N5.18 appears as a good candidate to elicit plant metabolism and consequently, to increase productivity of Papaver somniferum.
8. Fairbairn, J. W. and {El-Masry}, S., The Alkaloids ofPapaver Somniferum L.—VI.: “Bound” Morphine and Seed Development, Phytochemistry, vol. 7, no. 2, pp. 181--187, February 1968. doi: 10.1016/S0031-9422(00)86313-X. Abstract
   Vigorous acid hydrolysis or pepsin digestion of ground poppy seeds led to the production of alkaloid-like substances including codeine. Fermentation of ground seeds or germination of whole seeds for 1 to 2 days also led to the production of similar compounds as well as papaverine. Radioactive morphine was fed to the latex of the capsule and the seeds collected later were germinated in similar conditions and the alkaloid-like substances shown to be radioactive. Furthermore the ovules were shown to contain radioactive alkaloid-like substances (without degradative treatment) as well as radioactive non-alkaloidal compounds similar to those already reported. It is suggested that morphine is rapidly metabolised in the latex into a series of compounds some of which are alkaloid-like and others non-alkaloidal, ethanol insoluble, “bound” forms. The bound forms are stored in the seeds and broken down into smaller alkaloid-like substances during germination. An attempt was made to produce seeds deficient in these compounds by depriving the capsule of much of its latex at a critical stage in development. The seeds produced showed remarkable differences from controls; their germination rate and root hair and chlorophyll production were significantly reduced, thus indicating that the latex contents are essential to the production of viable seeds. The possible pharmacological effect of bound forms of alkaloids in poppy seeds is also discussed.
9. Fernández, Alfonso and Sáiz, Francisco, (Oryctolagus Cuniculus L.) as Seed Disperser of the Invasive Opium Poppy (Papaver Somniferum L.), Mastozoología Neotropical, vol. 14, no. 1, pp. 19--27, 2007. url: https://www.redalyc.org/pdf/457/45714103.pdf. Abstract
   We investigated whether introduced European rabbit on Robinson Crusoe Island, Chile, has the potential to spread the seeds of the exotic Papaver somniferum (opium poppy) via endozoochory. A total of 1320 rabbit droppings were collected during early autumn in the study area and analyzed in laboratory with binocular magnifying glass. Seeds from six different species were found in the droppings: Papaver somniferum (Papaveraceae), Centaurea melitensis (Asteraceae), Amaranthus sp. (Amarantaceae), Melilotus indicus (Fabaceae), Rumex acetosella (Polygonaceae), and one unidentified species. Poppy seeds were dominant. Most seeds (82\%) were destroyed during ingestion, but some of the intact seeds found in pellets remained viable. Germination rate of ingested seeds was similar to control seeds but germination success was lower in the former. Poppy seeds were found in pellets deposited inside poppy patches but also up to 100 m away from patches. In this way, rabbit expands the seed shadow of the plant. Our results suggest that, although poppy lacks morphological adaptations to long distance dispersal, seeds reach favorable places for seedling germination and recruitment via consumption by rabbits, an unspecialized seed disperser. Rabbit activity, together with the effect of other animals and some abiotic factors—wind and rain water, principally—contributes to the spread of poppy with implications for the management of both invasive species in the island.
10. Grover, I.S. and Dhanju, M.S., Effect of Gamma Radiation on the Germination of Papaver Somniferum and P Rhoeas, Indian Journal of Plant Physiology, vol. 22, no. 1, pp. 75--77, 1979. Abstract
    The germination of opium poppy (P somniferum) was increased whereas those of red poppy (Prhoeas) decreased considerably The differential radiosensitivity may be the cause of different result obtained in these two species (auth)
11. Hakimi, Yousef and Graduate, M and Fatahi, Reza and Naghavi, Mohammad Reza and Zamani, Zabihollah, Seed Germination Indices of Papaver Bracteatum Populations under Different Temperature Treatments, vol. 5, October 2021. url: https://www.researchgate.net/publication/361408878_Seed_germination_indices_of_Papaver_bracteatum_populations_under_different_temperature_treatments. Abstract
    Medicinal plants are sources of active and valuable secondary metabolites that have been economically beneficial for pharmacy and medicine. However, it is challenging to start large-scale and commercial cultivation of these plants because most of the arable land is mainly used to produce strategically essential crops. Persian poppy (Papaver bracteatum Lindl.) is a perennial medicinal plant belonging to the Papaveraceae family endemic to the mountainous areas in Northern Iran. It is known for high amounts of the valuable benzylisoquinoline alkaloid thebaine. This study investigated the effect of temperature treatments on germination characteristics of different Papaver braceatum populations on germination characteristics. For this purpose, three-level of temperature on three different populations of Persian poppies. The interactions of the population×temperature was significant for germination characteristics. The results of this experiment generally showed a decrease in all germination characteristics, including germination percentage, germination rate, plumule length, radical length, plumule dry weight, radical dry weight, and seed vigor index with increasing temperature level. The highest germination percentage in all three populations was recorded at 25 ° C, and the highest seed vigor index was recorded at 20 ° C. According to these results, to industrially cultivate this plant, the temperature is the most critical factor for achieving maximum yield. Therefore, to maintain the balance between vegetative growth and increase in plant dry weight, it is recommended to set the temperature in the range of 20 to 25 ° C because it was observed that the highest vegetative growth was recorded at 20 ° C and the highest dry weight at 25 ° C.
12. Karlsson, L. M. and Milberg, P., A Comparative Study of Germination Ecology of Four Papaver Taxa, Annals of Botany, vol. 99, no. 5, pp. 935--946, May 2007. doi: 10.1093/aob/mcm045.
13. Karlsson, L. M. and Milberg, P., Seed Dormancy Pattern and Germination Preferences of the South African Annual Papaver Aculeatum, South African Journal of Botany, vol. 73, no. 3, pp. 422--428, July 2007. doi: 10.1016/j.sajb.2007.03.007. Abstract
    Seeds from two populations of Papaver aculeatum, collected in South Africa, were subjected to a series of experiments to investigate their dormancy and germination. Dormancy was weak: fresh seeds germinated to nearly 100\% at 20/10 and 25/15~°C day/night if provided with light, up to 50\% at 15/5, but not at all at 30/20~°C. Warm stratification increased germination, both in darkness and at 15/5~°C, but did not lead to germination at 30/20~°C. Cold stratification reduced germination and limited germination to the cooler temperatures. Alternating cold and warm stratifications showed that the species undergoes dormancy cycles. When subjected to three different artificial annual changing climates, with onset both in summer and autumn, most seed germinated in the first autumn. However, in cooler climates, some germination occurred during spring and the second autumn. The general dormancy pattern, reduction during a warm period and induction during a cold, was similar to that of Papaver rhoeas, a European species and a wide-spread weed. However, germination temperature preferences differed by P. aculeatum avoiding germination at high temperatures, having substantially weaker dormancy when fresh and having more easily reduced dormancy, compared with P. rhoeas. These differences most likely result in differing temporal pattern of germination in the field.
14. Khan, Bilal Ahmad and Nadeem, Muhammad Ather and Afzal, Sadia and Afzal, Sadia and Khan, Muhammad Azim and Abbas, Tasawar and Aziz, Ahsan and Hasan, Syed Wasim and Amin, Muhammad Mohsin and Aziz, Amir and Nazeer, Safdar, Allelopathic Influence of Poppy (Papaver Somniferum L.) on Emergence and Initial Seedling Growth of Red Rice (Oryza Punctata L.), JOURNAL OF WEED SCIENCE RESEARCH, vol. 27, no. 2, pp. 381--392, December 2020. doi: 10.28941/pjwsr.v26i4.850. Abstract
    Allelopathy plays crucial role in effective weed control. Opium (Papaver somniferum L.) crop release different allelochemicals at maturity which have potential to act as natural weed killer in different crops. Phytotoxic effect of poppy (P. somniferum) was examined on emergence and initial seedling growth of red rice (Oryza punctate L.). Aqueous extract of different plant parts (leaves, stem and flower) was used at various concentrations (0.25\%, 0.5\%, 1\%, 2\%, 4\% and 8\%) along with distilled water as control. The aqueous extracts of leaves, stem and flower of P. somniferum significantly inhibited the emergence, seedling growth as well as root length (cm), shoot length (cm), fresh weight (g) and dry weight (g) of O. punctata. Maximum mean emergence time (9.18 days) and minimum shoot length (1.13 cm) emergence index (0.89) and emergence percentage (6.67\%) was observed under fruit extract at 8\% concentration. P. somniferum aqueous extract of stem at 8\% concentration took maximum time to complete 50\% emergence and gave minimum root length, fresh weight, and dry weight of O. punctata. Based on these finding it can be concluded that the phyto-chemicals present in P. somniferum can be used as eco-friendly O. punctata growth inhibitor to manage this weed in crops especially under organic cropping.
15. {Muntaha Tul Sidra} and {Sayeda Sarah Mubarak} and {Fatima Khan Sherwani} and {Nazma Azeem}, Allelopathic Potential of Papaver Somniferum L. on Seed Germination Against Three Different Varities of Zea Mays, February 2021. doi: 10.21203/rs.3.rs-181838/v1. Abstract
    Aims To evaluate the allelopathic effect of Papaver somniferum against the 3 varieties of Zea mays seeds cv. Azam, Pahari and Iqbal.Methods Aqueous extracts were made by soaking the powder of dried leaves of Papaver somniferum. For the aqueou...
16. Ograjšek, Simon and Ačko, Darja KOCJAN and Kastelec, Damijana, The Impact of the Period of Sowing and Fertilization on Morphological Characteristics and Seed Yield of Garden Poppy (Papaver Somniferum L.), Acta agriculturae Slovenica, vol. 107, no. 1, pp. 219--228, April 2016. doi: 10.14720/aas.2016.107.1.20. Abstract
    Garden poppy (Papaver somniferum L.) is a traditional crop that had already been cultivated in Slovenia in the past for the production of seed and oil. During its re-introduction in present time, numerous agro-technical dilemmas have been raised for the processes from sowing to harvesting and have to be studied in our present growing conditions. On the fields of the Ograjšek farm in Cerklje ob Krki two field experiments in complete randomized design had been sown in 2013 and 2014 to establish the influence of the sowing date and fertilization on morphological properties and crop yield of garden poppy seed of the Austrian variety 'Zeno 2002'. Results have shown that the average yield of seed in both trials (1643 kg/ha) is more than one time higher than the average crop yield recorded globally (700~kg/ha), proving how suitable growing conditions in the area of eastern Dolenjska are for the cultivation of poppy. Analysis of crop yield per plot showed there is no statistically significant interaction between the considered factors. The yield of poppy seed in the spring sowing term was 1742±77 kg/ha and in the autumn sowing was 1545±122 kg/ha. The difference is not statistically significant (p~=~0.1845). Recorded yield per plant was higher for the autumn term but that was not transferred to the total crop yield due to poor overwintering and consequentially lower plant density at the time of harvest. Both sowing dates have confirmed that exposure to light and duration of growing period had an important impact on plants height – average height of plants sown in autumn was 139±1 cm, which is more than 60 cm higher than the average height of the plants sown in spring (p = 0.0000). A strong statistical dependence of the yield of seed per capsule on diameter of the capsule was proved (p = 0.0000). It can be concluded that when the capsule diameter increases by 10~mm, the seed yield increases for 2.1 g to 2.3 g with 95 \% confidence. Dependence of the average seed yield on the number of lateral shoots per plant also proved to be statistically significant (p~=~0.0000). Linear model was used for the comparison of the four lines showed that the lines representing poppy sown in autumn and spring are statistically different for the control and for the ENTEC; the slope was higher for the poppy sown in autumn. These trials have given the first practical advice for production of garden poppy in Slovenia and should be continued for further useful results.
17. Rush, Michael D. and Kutchan, Toni M. and Coscia, Carmine J., Correlation of the Appearance of Morphinan Alkaloids and Laticifer Cells in Germinating Papaver Bracteatum Seedlings, Plant Cell Reports, vol. 4, no. 5, pp. 237--240, October 1985. doi: 10.1007/BF00269366. Abstract
    The course of alkaloid accumulation and laticifer cell appearance was compared in germinating P. bracteatum seedlings. Seedlings of various ages (0–14 days old) were analyzed for their dopamine, thebaine, morphinan alkaloid immunoreactivity, and benzophenanthridine alkaloid levels. Simultaneous electron microscopic studies revealed that seedlings were devoid of laticifer initials until day 3, where-upon their numbers increased with time. The appearance of appreciable amounts of thebaine only occurred after day 4 of germination. Conversely, dopamine was rapidly formed at the onset of germination and reached millimolar concentrations well before laticifer cells were detected. Benzophenanthridine alkaloid levels remained fairly constant over the period analyzed. These results support the theory that the presence of laticifer cells is necessary for the accumulation of morphinan but neither benzophenanthridine alkaloids nor their mutual precursor, dopamine.
18. Šerá, Božena and Novák, František, Stimulation of Seed Germination and Early Growth by Humic Substances on Poppy, Pepper, Rape, and Hemp, Biologia, vol. 77, no. 3, pp. 641--648, March 2022. doi: 10.1007/s11756-021-00952-1. Abstract
    Stimulatory effect of humic and fulvic acids (humic substances) extracted from local sources of lignite and soils on germination and early growth of hemp (Cannabis sativa L.), poppy (Papaver somniferum L.), pepper (Capsicum annuum L.) and rape (Brassica napus subsp. napus) were verified. The following structural parameters were determined for humic substances: aromaticity, biological activity, and hydrophilicity/hydrophobicity ratio. Germination rate, weight of seedling, accretion of seedling, and seedling vitality were determined in a growth test in laboratory conditions. It was found that the type of plant together with the type of humic and fulvic acids was of great importance for germination and initial growth (two-way ANOVA). The reaction of the monitored plant species was different to the types of humic and fulvic acids. A positive effect on germination and initial growth was observed in poppies (compared to controls, p\,{$<$}\,0.05); in other species, no significant statistical responses were observed. The tested plants can be sorted according to the descending significant positive effect on the weight of seedling as follows: poppy, pepper, rape, hemp. We found that structural parameters of humic and fulvic acids correlated with many measured parameters of germination and early growth (p\,{$<$}\,0.05). The results show that the origin of humic substances and their structural properties can significantly affect the germination of seeds and their initial growth. In addition, the responses of taxonomically different plant species after treatment with humic acids can be fundamentally different.
19. Singh, S. and Singh, A.K. and Kumar, B., Effect of Moisture Regimes and Field Practices on Germination and Establishment of Opium Poppy (Papaver Somniferum L.), Acta Horticulturae, no. 1036, pp. 169--173, May 2014. doi: 10.17660/ActaHortic.2014.1036.19. Abstract
    Opium poppy (Papaver somniferum L.) is an important plant grown for opium extraction from its capsules. At present legal cultivation of opium poppy is going on in the three adjoining states of India viz. Uttar Pradesh, Madhya Pradesh and Rajasthan. In general, the germination of seeds of this crop is very good (80-95\%) under laboratory conditions. Germination is poor under field conditions and farmers use higher quantity of seeds to establish normal plant populations. Therefore, there is a need to develop ideal field practices to improve the germination and reduce seed rate under commercial cultivation. A field experiment was conducted in November 2007 to evaluate the effect of two moisture regimes: sowing of opium poppy seed at 60\% available soil moisture after applying presowing irrigation (I1) and sowing of opium poppy seeds in dry seed bed followed by irrigation (I2) with five field practices: sowing of seeds on soil surface (P1), sowing of seeds in 2.5 cm deep furrows and covered with soil (P2), sowing of seed in 2.5 cm deep furrows and kept open (P3), sowing of seeds in 5.0 cm deep furrows and covered with soil (P4) and sowing of seed in 5.0 cm deep furrows and kept open(P5). The seeds of cultivar ‘Sampada’ having 94 and 57\% germination under laboratory and earthen pot conditions, respectively were used for these studies. The soil of experimental site was sandy loam in texture. As a result of the field experiment maximum germination percentage and plant stand (59\%) was recorded in treatment combination I1P2. The next best treatment with respect of germination percent and plant stand was I2P1 where germination was recorded as 35\%. Results of these studies suggest that opium poppy should be shown at 60\% available soil moisture in 2.5 cm deep furrows and furrows should be covered with soil in order to obtain higher germination rates and plant stand under field condition.
20. Kara, Nimet and Baydar, Hasan, The Influence of Sowing and Planting Seedlings at Different Dates in Autumn on the Yield and Quality of the Opium Poppy (Papaver Somniferum L.), Journal of Applied Research on Medicinal and Aromatic Plants, vol. 21, pp. 100290, March 2021. doi: 10.1016/j.jarmap.2020.100290. Abstract
    Turkey is the world’s leading country in legal poppy cultivation and morphine production. As with other producing countries, the poppy is traditionally cultivated in Turkey through the sowing of seeds during the autumn. However, if the seasonal rains are insufficient and irregular, this can lead to problems with germination and emergence. In this research, the option of solving these problems by using seedlings in production, instead of seeds, was investigated. The study compares the yield and quality performance of the two production methods, planting seedlings and direct seed sowing in the poppy cultivar ‘Ofis-4’. Field experiments were conducted on three sowing/planting dates, namely October fifteenth, November first, and November fifteenth, during the growing seasons of 2017/18 and 2018/19 in the semi-arid ecological conditions of the southwestern region of Turkey. The plants produced by seedling transplantation and by sowing were compared in terms of the yields of seeds and capsules along with yield-related traits and the contents of seed-fixed oil and capsule opium alkaloids. The planting seedling method was seen to be an average of one week faster than the sowing method. In terms of the sowing/planting time delays in the autumn season, it was also observed that the ripening difference between both propagation methods started to narrow. If the general averages are considered, the highest capsule and seed yields were obtained from planting seedling on October fifteenth, the first planting date, in both years. Since the planting dates were late, the capsule and seed yield decreased while it was found that the contents of capsule alkaloids, except morphine and codeine, were not significantly affected by the production method and sowing/planting dates. On the other hand, the direct sowing method and first sowing date produced the highest morphine (0.53 and 0.58 \%, respectively) and codeine (0.101 and 0.126 \%, respectively) contents. It was therefore concluded that the seedling planting method can be recommended as an alternative to the direct sowing method traditionally applied in poppy production for the achieving of high capsule and seed yields.
21. Stranska, Irena and Skalicky, Milan and Novak, Jan and Matyasova, Elena and Hejnak, Vaclav, Analysis of Selected Poppy (Papaver Somniferum L.) Cultivars: Pharmaceutically Important Alkaloids, Industrial Crops and Products, vol. 41, pp. 120--126, January 2013. doi: 10.1016/j.indcrop.2012.04.018. Abstract
    The content of pharmaceutically important alkaloids (morphine, codeine, thebaine, papaverine and narcotine) was analysed in the empty poppy capsules (poppy straw) of fifteen selected cultivars of Papaver somniferum. By applying the cluster analysis and k-means methods, four clusters representing the sets of cultivars with a defined level of production of the selected alkaloids were determined, based on their similarities. The content of individual minor alkaloids, which was obtained was relatively low (0.00–0.17\%) and in some cases negligible, defined by the dominant representation of morphine in the alkaloid spectrum of all selected cultivars. In addition to the high-morphine cultivar ‘Buddha’ (mean 1.64\%), there was a middle content (mean 0.79\%) or low content (mean 0.61\%) of morphine in the cultivars included in clusters 2 and 3 (average of 3 consecutive years 2007, 2008 and 2009). A relatively high content of morphine was detected in cultivar ‘Böhmův bělosemenný’ (mean 0.91\%), although it is usually published that white-seed cultivars contain less morphine and other alkaloids, compared to blue-seed or grey-seed cultivars. Low content of morphine (mean 0.48\% resp. 0.44\%) by relatively high content of narcotine (mean 0.11\% resp. 0.15\%) was obtained in poppy cultivars ‘Danneborg’ and ‘Tatarstan’. Also, surprisingly the highest papaverine content (mean 0.07\%) was detected in the cultivar ‘Sušický červenosemenný’ with the lowest content of morphine (mean 0.34\%).
22. Yazici, Levent, Influence of Different Sowing Times on Yield and Biochemical Characteristics of Different Opium Poppy (Papaver Somniferum L.) Genotypes, Journal of King Saud University - Science, vol. 34, no. 8, pp. 102337, November 2022. doi: 10.1016/j.jksus.2022.102337. Abstract
    Background Opium poppy (Papaver somniferum L.) is an important industrial plant that could be cultivated in Turkey during autumn and spring seasons. However, the impact of different seasons on yield and biochemical attributes of different genotypes in Turkey is rarely tested. This study determined the effect of different sowing times (autumn and spring) on alkaloid, oil content, yield and related traits, and phenological properties of different opium poppy genotypes. Methods The experiment was conducted at two different locations (Topcu and Yerkoy) during autumn and spring seasons of 2020 and 2021. A total 18 different poppy genotypes available for cultivation were included in the study. Data on yield related traits and biochemical attributes were recorded at harvest. Results Alkaloid and oil contents were higher in spring sowings than autumn sowings. However, autumn sowings recorded higher yield and yield-related traits compared to spring sowings. During autumn sowings, morphine ratio varied from 0.33~\% to 1.40~\%, morphine yield ranged between 4.28 and 21.97~kg~ha−1, capsule yield differed between 1156.1 and 2393.3~kg~ha−1, and seed yield ranged from 1162.0 to 2420.0~kg~ha−1. Similarly, during spring sowings, morphine ratio differed from 0.40~\% and 1.50~\%, morphine yields ranged between 3.13 and 17.44~kg~ha−1, capsule yield varied between 721.7 and 1757.8~kg~ha−1, and seed yield ranged from 656.0 to 1676.4~kg~ha−1. The genotypes ‘Celikoglu’, ‘Tinaztepe’, ‘Seyitgazi’, ‘Ofis 95’ and ‘Afyon 95’ are recommended for autumn sowing, whereas genotypes ‘Huseyinbey’, ‘Ofis 96’, ‘Celikoglu’, ‘Seyitgazi’ and ‘Ofis NM’ are recommended for spring sowing due to their higher yield and yield characteristics. The genotypes ‘Ofis 1’ and ‘Ofis 2’ can be used in both autumn and spring sowings for morphine production. The genotypes ‘Ofis 8’ and ‘TMO 1’ during autumn sowings, and ‘Ofis 96’ and ‘TMO 1’ are recommended for spring sowings to obtain higher oil contents. Conclusion The results have grouped and identified the available genotypes according to their specific traits. Therefore, the selection of the genotype must be done for the required purpose based on the results of the current study.
23. Luhmer, Katharina and {Schulze-Kaysers}, Nadine and Feuereisen, Michelle and Wirth, Lukas and Maretzky, Fabian and Wüst, Matthias and Blum, Hanna and Dörr, Elisa and Pude, Ralf, Fatty Acid Composition, Tocopherols, Volatile Compounds, and Sensory Evaluation of Low Morphine Yielding Varieties of Poppy (Papaver Somniferum L.) Seeds and Oils, Journal of Agricultural and Food Chemistry, vol. 69, no. 11, pp. 3439--3451, March 2021. doi: 10.1021/acs.jafc.0c07183. Abstract
    Low morphine yielding winter (‘Zeno Morphex’) and summer (‘Viola’, ‘Mieszko’, ‘Borowski’) poppy varieties were investigated for their chemical composition and sensory properties. The oil content of the 13 seed samples as well as that of fatty acids, tocopherols, and volatile compounds in the respective oils were determined, and the sensory profiles of the seeds and oils were established. Linoleic acid made up 70.7–75.2\% of the fatty acids. High amounts of γ-tocopherol were detected, especially in variety ‘Viola’ (287 ± 34 mg kg–1), while δ-tocopherol was only present in ‘Zeno Morphex’ (3.9 ± 0.6 mg kg–1). The most abundant volatiles were caproic acid (1.4–148 μg g–1), hexanal (0.9–15.2 μg g–1), 1-hexanol (0.3–20.1 μg g–1), limonene (1.3–9.4 μg g–1), and 2-pentylfuran (1.0–7.8 μg g–1). The sensory panel distinguished samples in particular by fatty/oily, rancid, sweet, and green attributes, the green aroma being correlated to three methoxypyrazines only present in summer poppies.
24. Kara, Nimet, The Effects of Autumn and Spring Sowing on Yield, Oil and Morphine Contents in the Turkish Poppy (Papaver Somniferum L.) Cultivars, Turkish Journal Of Field Crops, vol. 22, no. 1, pp. 39--46, March 2017. doi: 10.17557/tjfc.301829. Abstract
    The poppy is a plant that can be planted in the autumn and spring seasons. The purpose of the research was to investigate the effects of autumn and spring sowing on capsule yield, seed yield, yield components, oil content and morphine content of fifteen poppy cultivars that have been recently registered under Isparta conditions. The experiment was arranged according to the randomized complete-block design with three replicates during the years of 2012/13 and 2013/14. The results of the research showed that the differences among capsule yields, seed yields, yield components and morphine contents of the cultivars were statistically significant in both sowing times and years. The highest and the lowest values of plant height, capsules number, capsule width, capsule length, 1000 seed weight and oil yield of the poppy cultivars varied according to the cultivars, the sowing times and the years. The capsule yield and seed yield of the poppy cultivars in the autumn sowing varied between 416.7-1043.3 kg ha-1 and 523.5-1276.3 kg ha-1 in the first year, between 465.3-1375.6 kg ha-1 and 596.7-1520.4 kg ha-1 in the second year, respectively. In the spring sowing, the capsule yield and seed yield of the poppy cultivars varied between 204.3-455.7 kg ha-1 and 286.0-573.0 kg ha-1 in the first year, between 497.2-830.7 kg ha-1 and 614.3-962.7 kg ha-1 in the second year, respectively. The highest oil content was determined in the Zaferyolu cultivar in both years and sowing times. The morphine content of the poppy cultivars varied between 0.47-1.00\% in the autumn sowing and between 0.45-0.97\% in the spring sowing, respectively. The higher values in the autumn sowing were obtained than the spring sowing in all of the characteristics investigated in both years, except for the oil content. Considering the present results, TMO-3, Ofis-4, Ofis-8, Tınaztepe and Zaferyolu cultivars were advised in the autumn sowing and the TMO-1, Ofis-8 and Afyon-95 cultivars in the spring sowing because of their higher capsule, seed and oil yields. In point of the morphine contents, the TMO-3, TMO-T, Ofis-3, Ofis-8 and Bolvadin-95 cultivars were advised.
25. Solanki, Gurudatt and Dodiya, N.S. and Kunwar, Ram and Bisen, Prashant and Kumar, Ramesh and Singh, Jagjeet and {dhya}, San, Character Association and Path Coefficient Analysis for Seed Yield and Latex Yield in Opium Poppy (Papaver Somniferum L.), International Journal of Current Microbiology and Applied Sciences, vol. 6, no. 8, pp. 1116--1123, August 2017. doi: 10.20546/ijcmas.2017.608.138.
26. Singh, S. P. and Shukla, Sudhir, Stability Parameters for Opium and Seed Yield in Opium Poppy (Papaver Somniferum), Indian Journal of Agricultural Sciences, vol. 71, no. 5, pp. 313--315, May 2001. url: http://epubs.icar.org.in/ejournal/index.php/IJAgS/article/view/39797. Abstract
    An experiment was conducted during 1995-96 to 1998-99 to evaluate 10 varieties for stability parameters for opium yield, seed yield and morphine content in opium poppy (Papaver somniferum L.). The varieties showed significant differences for all the traits. Joint regression analysis indicated that both predictable and unpredictable components contributed significantly towards the difference in stability of genotypes. However, larger variation in regression coefficients revealed that genotypes had different degree of environmental response. Genotypes studied did not show uniform stability (s2 di) and response (linear) for all the characters. The genotypes 'NBRI 9', 'NBRI 10' and 'BROP 1' were more stable and responsive than other over environment for all 3 traits. The mean performance was found associated with linear (S2 di) component with mean (X) and regression (bi) was noticed.
27. Mirjana, J. and Sonja, I. and Gjoshe, S. and Jelena, A. and Ljupcho, J. and Dane, B., Interrelationship between Seed and Capsule Yield and Yield Contributing Characters in Opium Poppy (Papaver Somniferum L.)., Agrarni Nauki, vol. 4, no. 11, pp. 13--17, 2012. url: https://www.cabdirect.org/cabdirect/abstract/20133050876. Abstract
    The aim of this study was to obtain information on the association of different characters in a collection of 49 opium poppy genotypes, as well as to determine the traits that have the highest direct effect on the seed and capsule yield. Significant positive correlations were observed between the seed yield and plant height, number of capsules per plant, diameter of main capsule, weight of main...
28. {Mahdavi-Damghani}, Abdolmajid and Kamkar, Behnam and {Al-Ahmadi}, Majid Jami and Testi, Luca and {Muñoz-Ledesma}, Francisco J. and Villalobos, Francisco J., Water Stress Effects on Growth, Development and Yield of Opium Poppy (Papaver Somniferum L.), Agricultural Water Management, vol. 97, no. 10, pp. 1582--1590, October 2010. doi: 10.1016/j.agwat.2010.05.011. Abstract
    The effects of pre-anthesis water deficit and cycle length were examined in Papaver somniferum L., cultivated for alkaloid production, in two locations in southern Spain. The vegetative period was shortened by extending the photoperiod through supplemental lighting in the field, while water deficit in pre-anthesis was induced by avoiding irrigations and installing rain shelters. The treatments were: IN (irrigated-normal photoperiod), IL (irrigated-hastened flowering), DN (water deficit in pre-anthesis-normal photoperiod) and DL (water deficit in pre-anthesis and hastened flowering). The artificial photoperiod hastened the flowering by 15 and 21 days, for irrigated and deficit treatments respectively. Seasonal evapotranspiration (ET) ranged from 398 (DN) to 505mm (IN). There was evidence of root water uptake deeper than 1.5m. Stomatal conductance was reduced (16\%) during water stress, and did not recover in post-anthesis after resuming irrigation. Head yields (capsule+seeds+7cm stem) ranged between 3.8 and 4.3tha−1; water deficit and short vegetative period both reduced the biomass accumulated, although the effect on yields in these treatments was counterbalanced by a higher harvest index. Early flowering had a detrimental effect on alkaloid concentration in the capsule. Alkaloids yield ranged between 27 and 37kgha−1. Water use efficiency (WUE) ranged between 0.78 and 0.96kgm−3 ET for yield and between 63.4 and 73.7gm−3 ET for alkaloids. Water stress increased slightly the Water Use Efficiency. A shorter vegetative phase had no effect on WUE for biomass or yield, but decreased the WUE for alkaloids production.
29. Lošák, Tomáš and Elbl, Jakub and Kintl, Antonín and Ducsay, Ladislav and Varga, Ladislav and Árvay, Július and Neugschwandtner, Reinhard W. and Vollmann, Johann and Gondek, Krzysztof and {Mierzwa-Hersztek}, Monika and Demková, Lenka and Čermák, Pavel, Effect of Poppy (Papaver Somniferum L.) Fertilization with Potassium and Magnesium on the Seed Yield and Its Quality, Acta fytotechnica et zootechnica, vol. 25, no. 3, pp. 242--246, October 2022. doi: 10.15414/afz.2022.25.03.242-246. Abstract
    In a vegetation pot experiment we explored the effect of fertilizers with potassium (KCl; 0.9 g K2O/6 kg of soil) and magnesium (ESTA Kieserite; 0.3 g MgO/6 kg of soil) applied together or separately to poppy on soil with good K and Mg supply on the seed yield, oil content and cadmium in the seed. Seed yields increased significantly after the application of KCl and ESTA Kieserite, respectively by 23.89\% and 21.68\%, as compared to the unfertilized control. Combined application of both fertilizers stimulated the yields significantly, i.e. by 15.26\%. The seed oil content showed no significant changes among the treatments and ranged only between 42.1 and 43.6\%. The pot production of oil in the fertilized treatments increased to 2.21-2.44 g/pot compared to 1.92 g/pot in the unfertilized control. The Cd content in seeds was very low, there were no differences among the treatments and it ranged only between 0.204 and 0.214 mg/kg fresh matter - FM (threshold level of 1.20 mg/kg wet weight). Therefore, the application of both KCl and ESTA Kieserite fertilizers can be recommended on soils well supplied with both nutrients.
30. Bernáth, J. and Németh, É. and Petheõ, F. and Friedt, W., Alkaloid Accumulation in Capsules of the Selfed and Cross-Pollinated Poppy, Plant Breeding, vol. 122, no. 3, pp. 263--267, 2003. doi: 10.1046/j.1439-0523.2003.00811.x. Abstract
    In harmony with international regulations the creation of poppy cultivars with high alkaloid content as well as free of alkaloids is in progress. In the course of selection procedures, five poppy cultivars of high chemical diversity, ‘Kheops’, ‘AT, ‘Tebona’, ‘Kék Gemona’ and ‘Przemko’, were selfed and crossed in two successive years (2000-01). After full ripening, both the biomass production and the alkaloid content of capsules of selfed and crossed plants were determined. When the castrated alkaloid-free cultivar ‘Przemko’ (accumulating only 0.01 mg/100 g morphine in capsules) was pollinated with cultivars rich in alkaloids, the morphine content of its capsules increased to 0.9-7.5 mg/100 g-values. Based on the well-known biosynthetic background of poppy alkaloids, this can be explained by promotion of (S)-norcoclaurine synthesis, which seems to be suppressed in alkaloid-free plants; but tissues of the developing hybrid seeds, which contain a combined gene pool of both alkaloid-free cultivar and that of the pollen donor plants, release this suppression. The chemical metaxenia also manifested itself in other combinations. In particular, the appearance of narcotine (0.1-1.1 mg/100 g) in the narcotine-free cultivars ‘Kheops’, ‘A1’ and ‘Przemko’ can be explained biochemically. If the narcotine-type plants ‘Kék Gemona’ (accumulating 11.3 mg/g narcotine) were used for pollination, the developing seed tissue, which contains the gene pool of narcotine biosynthesis, might contribute to the suppression of the activity of 1,2-dehydroreticuline reductase, which leads to the accumulation of narcotine at the cost of morphinanes.
31. Meos, A and Saks, L and Raal, A, Content of Alkaloids in Ornamental Papaver Somniferum L. Cultivars Growing in Estonia, Proceedings of the Estonian Academy of Sciences, vol. 66, no. 1, pp. 34, 2017. doi: 10.3176/proc.2017.1.04. Abstract
    The alkaloid content in industrial opium poppies (Papaver somniferum L.) has been studied by numerous authors, but not in ornamental cultivars.The aim of the study was to investigate the content of morphine, codeine, and papaverine in P. somniferum cultivars by high performance liquid chromatography (HPLC) and capillary electrophoresis (CE). The studied poppy capsules were either grown from the ornamental poppy seeds purchased from different vendors or were unknown cultivars from home gardens in Estonia, cultivated as ornamental poppies. All dried capsules samples (n = 34) contained on average 362 mg/100g of morphine, 25 mg/100g of codeine, and 29 mg/100g of papaverine as determined by CE. The content of morphine determined by HPLC was 152–676 mg/100g, which is a remarkably high concentration allowing potential illegal use of ornamental poppies. Both analytical methods demonstrated similar results. Hence, CE is a suitable alternative to HPLC for quantitative analysis of opium alkaloids as it simplifies the sample preparation procedure.
32. Singh, M. and Chaturvedi, N. and Shasany, A.K. and Shukla, A.K., Impact of Promising Genotypes of Papaver Somniferum L. Developed for Beneficial Uses, Acta Horticulturae, no. 1036, pp. 29--41, May 2014. doi: 10.17660/ActaHortic.2014.1036.3. Abstract
    Papaver somniferum (opium poppy) is a treasure medicinal plant known to produce therapeutically useful alkaloids like morphine, codeine, thebaine, papaverine, etc. Its seeds have high nutritional value (oil is rich in unsaturated fatty acids). Morphine, which is used as an indispensable analgesic, is in high demand. However, the narcotic nature of morphine and its easy illicit conversion into heroin has curtailed the free agriculture of P. somniferum and restricted the benefits that may accrue to the poppy farmers. Since the poppy alkaloids cannot be chemically synthesized in an economically viable manner, the plant remains the sole source of the biologically active alkaloids. Most of the licit morphine is chemically converted to codeine, which has a high pharmaceutical demand. In recent years, the trend is shifting towards thebaine and oripavine (which are precursors in the in planta morphine biosynthetic pathway) derived drugs as it is becoming increasingly difficult to control the narcotic usage of morphine. This has led to a global demand for poppy cultivars that are either (i) low in morphine but produce substantial amounts of thebaine, oripavine and codeine or (ii) are latex-less so that upon lancing the capsule do not exude latex (that contains morphine). Opium poppy crop is plagued by many devastating diseases (e.g. downy mildew). Therefore, there is a need to develop/identify disease-resistant genotypes. Significant efforts have been made globally to achieve these objectives and several speciality genotypes/cultivars of opium poppy are presently available. This paper discusses various aspects of the opium poppy crop including the present status and bottlenecks. It also highlights the future course of action to be taken for research on opium poppy.
33. Gümüşçü, Ahmet and Arslan, Neşet and Sarıhan, Ercüment Osman, Evaluation of Selected Poppy (Papaver Somniferum L.) Lines by Their Morphine and Other Alkaloids Contents, European Food Research and Technology, vol. 226, no. 5, pp. 1213--1220, March 2008. doi: 10.1007/s00217-007-0739-0. Abstract
    The alkaloid analysis was performed in the laboratory of the alkaloid factory located at Bolvadin province of Turkey in 2000 on selected 99 poppy lines, which were selected with respect to their morphological and agronomic characteristics from around 1,000 poppy lines provided by Agriculture faculty, Ankara University. Our results showed that the ratio of the alkaloids, morphine, thebaine, codeine, papaverine and noscapine ranged from 0.110 to 1.140\%, 0.005 to 0.134\%, 0.005 to 0.27\%, 0.001 to 0.440\% and 0.006 to 0.418\%, respectively, in the selected 99 poppy lines. According to their alkaloid content, 15 lines had significantly higher morphine, three lines had higher noscapine, and one line had higher papaverine levels.
34. Kumar, Birendra and Patra, N. K., Gene Frequency-Based Estimation of Natural Outcrossing in Opium Poppy (Papaver Somniferum L.), Molecular Breeding, vol. 26, no. 4, pp. 619--626, December 2010. doi: 10.1007/s11032-010-9395-3. Abstract
    Mating systems are generally thought to be important factors in determining the amount and nature of genetic variability in a population. Nearly 1,000 individuals at a single location (Lucknow) and over two~years were crossed and subsequently scored for selfing versus outcrossing in 9–10 monohybrid populations of opium poppy (Papaver somniferum). Three different alleles of two marker loci (two—R/r and P/p—for anthocyanin locus and B/b for capsule size locus) were used to determine the male gametes that had effected fertilizations in F2 recessives (rr, pp and bb). The estimates of the gene frequency-based outcrossing parameter (α) were found to vary with year, cross and marker locus used (α range: 7.21–71.03\%). Study of the two dihybrid crosses concerning the two marker loci simultaneously, further confirmed that outcrossing at the R/r or P/p locus was significantly greater than that at the B/b locus. The nature of the outcrossing was, in general, nonrandom. In this species, in general, selfing predominated, with one exception in respect of monohybrid crosses involving the purple form of anthocyanin locus, in which outcrossing predominated.
35. Yadav, H. K. and Shukla, Sudhir and Singh, S. P., Genetic Combining Ability Estimates in the F1 and F2 Generations for Yield, Its Component Traits and Alkaloid Content in Opium Poppy (Papaver Somniferum L.), Euphytica, vol. 168, no. 1, pp. 23--32, July 2009. doi: 10.1007/s10681-008-9872-5. Abstract
    The F1 and F2 generations of a twenty parent fractional diallel cross of opium poppy (P. somniferum L.) were analyzed for combining ability for ten quantitative and five quality (alkaloids) traits. The results indicated significant differences among the parents for combining ability for all the traits. The GCA and SCA components of variances were significant for all the characters. However, the SCA component of variance (δ2s) was predominant indicating the preponderance of non-additive gene effect for all the traits except for leaves/plant and papaverine in F1 hybrids. The average degree of dominance (δ2s/δ2g) was more than unity indicating over dominance and also confirming non-additive mode of gene action. Among the parents IS-16, IS-13 and NBRI-1 for early flowering, BR226 and BR241 for branches/plant, capsule weight/plant, seed yield/plant and husk yield/plant, BR227 for leaves/plant, UO1285 for capsule size and opium yield/plant, NBRI-5 for husk yield/plant, morphine, codeine, and thebaine and ‘Papline’ for plant height and papaverine content were found good general combiners. Parent ND1001 was good combiner for codeine and narcotine content. Inclusion of good general combiners in a multiple crossing program or an intermating population involving all the possible crosses among them subjected to bi-parental mating may be expected to offer maximum promise in breeding for higher opium, seed yield and alkaloid contents.