Resumen Introducción: Annona macroprophyllata Donn. Smith. (Annonaceae) es una especie de árbol frutal, conocida como papausa. En México y América Central, esta fruta se ha convertido en un cultivo importante y valorado debido a su delicioso sabor y la cremosa textura de su pulpa. Sus frutos son frecuentemente dañados por la incidencia de avispas del género Bephratelloides Girault (Hymenoptera: Eurytomidae), que se desarrollan dentro de las semillas. Objetivo: Describir la interacción de Bephratelloides cubensis Ashmead durante su ciclo de vida en frutos de A. macroprophyllata. Método: Recolectamos frutos en diferentes estados de crecimiento registrando a) la ovoposición, b) el momento de la infección evidente, c) el desarrollo de las avispas dentro de las semillas, y d) su emergencia como adultos. También determinamos la proporción de frutas y semillas dañadas. Resultados: Los datos indican que las avispas preferían ovipositar en frutas con un diámetro de menos de 8 cm, la oviposición era más frecuente entre las 11:00 a.m. y las 3:00 p.m., y que había un 26 % de la infestación en las frutas y un 9 % en semillas. Conclusión: Es una interacción obligada para la avispa y la mayor proporción de ataque a los frutos fue en las etapas tempranas de desarrollo de frutos y las acciones de control se deberían enfocar a la protección de estas etapas.

Abstract Introduction: Annona macroprophyllata Donn. Smith. (Annonaceae) (syn. Annona diversifolia Saff.) is a valued fruit tree species known as papausa. In Mexico and Central America, this fruit has become an important crop because of its tasty flavor and high pulp content. Its fruits are frequently damaged by the incidence of wasps of the genus Bephratelloides Girault (Hymenoptera: Eurytomidae), which develop inside the seeds. Objective: To report the interaction of Bephratelloides cubensis Ashmead during its life cycle in fruits of A. macroprophyllata. Methods: We periodically collected fruits in different states of growth recording a) oviposition, b) the moment of evident infection, c) the development of the wasps inside the seeds, and d) their emergence as adults. We also determined the proportion of damaged fruits and seeds. Results: The data indicate that wasps preferred to oviposit on fruits with a diameter of less than 8 cm, oviposition was more frequent between 11:00 am and 03:00 pm., and there was 26 % infestation of fruits, and 9 % of seeds. Conclusion: It is an obligatory interaction for the wasp, the highest proportion of attack on fruits was in the early stages of fruit development and control actions should focus on the protection of these early stages.

Abstract The genus Bursera is a characteristic component of the tropical dry forest (TDF) of Mexico, however, despite being relevant for the conservation and management of the species, its biology is little known, among them, their phenological patterns and their associations with seasonal factors. The aim of this research was to characterize the phenology of Bursera simaruba Jacq. ex L and Bursera tomentosa (Jacq.) Triana & Planch and to evaluate their relationship with environmental variables. The phenological study was realized in a protected area with tropical dry forest of the Central Depresion from Chiapas. Fifty adult trees of each specie were selected and the intensity of four reproductive phenophases (flower bud, open flower, fruit development and permanence of fruits) and three vegetative ones (leaf bud, mature leaves, senescence) were recorded using the Fournier method. Seasonality and synchrony in each phase were estimated by circular statistics calculating the mean vector (r) and the Rayleight test (Z). Also, the functional types of vegetative phenology were categorized and that of reproductive phenology. All the phenological stages were seasonal (r> 0.5), in both species, the flowering and leaf bud phenophases occurred in a short period while the vegetative and fruit production phenophases are the more extended stages. The reproductive and vegetative phenology of Bursera simaruba and B. tomentosa were very similar, and the most correlated environmental variables with them (rs ≥ 0.6) were temperature and humidity. These results allow to know the phenological dynamic of Bursera species in the TDF, and with this, to promote management and conservation strategies of these important species.

Resumen El género Bursera es un componente característico de los bosques tropicales secos (BTS) de México y relevante para la conservación de estos hábitats, sin embargo, sus aspectos fenológicos son poco conocidos. Este estudio tuvo como objetivo caracterizar la fenología de Bursera simaruba Jacq. ex L y Bursera tomentosa (Jacq.) Triana & Planch y evaluar su relación con las variables ambientales. El seguimiento fenológico se realizó en una zona protegida del BTS de la región florística conocida como Depresión Central en Chiapas, México durante un ciclo anual (mayo 2017- abril 2018). Se seleccionaron 50 individuos adultos de cada especie y se registró la intensidad de sus fenofases reproductivas (botón floral, flor abierta, desarrollo de frutos y permanencia de frutos) y fenofases vegetativas (brote foliar, hojas maduras, senescencia) utilizando los métodos de Fournier. La estacionalidad y sincronía de cada fase se estimó mediante estadística circular calculando el vector medio (r) y la prueba Rayleigh (Z). Además, se categorizaron los tipos funcionales de su fenología vegetativa y reproductiva. Las etapas fenológicas en ambas especies fueron estacionales (r > 0.5), la floración y brote foliar sucedieron en un periodo breve mientras que el desarrollo vegetativo y las de producción de frutos fueron etapas más extendidas. Las fenofases reproductiva y vegetativa de Bursera simaruba y B. tomentosa son similares, las variables ambientales más correlacionadas (rs ≥ 0.6) a ellas fueron la temperatura y la humedad. El estudio de estas burseras permite conocer su dinámica en el BTS y con esto proponer planes de manejo de estas especies resiníferas.

Resumo Estudos sobre o efeito da aplicação de reguladores vegetais para a superação da dormência de sementes têm sido realizados em espécies de Annonaceae, deste modo a relação endógena entre o ácido abscísico e as giberelinas, após a aplicação de reguladores vegetais, necessita ser mais bem estudada nesta família. Considerando a importância do balanço hormonal nos mecanismos de dormência, o objetivo deste trabalho foi elucidar como as concentrações endógenas de ABA e GA se modificam após a aplicação de reguladores vegetais em sementes durante a superação da dormência e como essas alterações afetam a degradação de reservas durante a germinação de sementes de Annona macroprophyllata e A. purpurea logo após tal superação. As sementes foram submetidas a três condições: - sem imersão; imersão em água, e imersão em GA4+7+benzyladenina. ABA, GA, lipídios, proteínas e açúcares solúveis totais foram quantificados aos 0; 2; 5; 10 e 15 dias após o início dos tratamentos. A aplicação dos reguladores levou à alteração do balanço hormonal, induzindo aumento do GA endógeno e redução dos níveis de ABA desde a embebição das sementes. Durante a embebição, foi observada a degradação de reservas das sementes (primeiro proteínas, depois os açúcares e finalmente os lipídios), e a germinação completa foi obtida após 10 dias do início dos tratamentos, com emissão da raiz primária. Em ambas as espécies, a dormência pode ser quebrada logo após a dispersão das sementes, quando o equilíbrio entre GA e ABA endógeno é alterado (pela aplicação exógena de reguladores vegetais), levando à degradação de reservas e à germinação.

Abstract Studies on the effect of the applying of phytoregulators to overcome seed dormancy have been carried out in Annonaceae species, thus the endogenous relationship between abscisic acid and gibberellins after the application of phytoregulators needs to be known in these species. Considering the importance of hormonal balance in dormancy mechanisms, the aim of this research was to elucidate how endogenous ABA and GA concentrations change after the application of phytoregulators in seeds and how these changes affect reserve degradation during germination of Annona macroprophyllata and A. purpurea seeds. Seeds were submitted to three conditions: - no soaking, soaking-in-water and soaking-in-GA4+7+benzyladenine. ABA, GA, lipids, proteins, and total soluble sugars were quantified at 0, 2, 5, 10, and 15 days after the beginning of treatments. The application of phytoregulators led change of the hormonal balance inducing increase in endogenous GA and reduction in ABA levels since seed soaking. During imbibition, the degradation of seed reserves (breaks proteins first, and then soluble sugars and finally lipids) was observed and complete germination was obtained after 10 days, with primary root emission. In both species, dormancy can be broken soon after seed dispersal when the balance between GA and ABA is endogenously changed (by exogenous application of phytoregulators), leading to metabolic reserve degradation and germination.

Abstract Background: Endemic systemic mycoses remain a health challenge, since these opportunistic diseases are increasingly infecting immunosuppressed patients. The simultaneous use of antifungal compounds and other drugs to treat infectious or non-infectious diseases has led to several interactions and undesirable effects. Thus, new antifungal compounds should be investigated. The present study aimed to evaluate the activity of liriodenine extracted from Annona macroprophyllata on agents of systemic mycoses, with emphasis on the genus Paracoccidioides. Methods: The minimum inhibitory concentration (MIC) and minimum fungicide concentration (MFC) were determined by the microdilution method. The cellular alterations caused by liriodenine on a standard P. brasiliensis (Pb18) strain were evaluated by transmission and scanning electron microscopy. Results: Liriodenine was effective only in 3 of the 8 strains of the genus Paracoccidioides and in the Histoplasma capsulatum strain, in a very low concentration (MIC of 1.95 μg.mL-1); on yeasts of Candida spp. (MIC of 125 to 250 μg.mL-1), including C. krusei (250 μg.mL-1), which has intrinsic resistance to fluconazole; and in Cryptococcus neoformans and Cryptococcus gattii (MIC of 62.5 μg.mL-1). However, liriodenine was not effective against Aspergillus fumigatus at the studied concentrations. Liriodenine exhibited fungicidal activity against all standard strains and clinical isolates that showed to be susceptible by in vitro tests. Electron microscopy revealed cytoplasmic alterations and damage to the cell wall of P. brasiliensis (Pb18). Conclusion: Our results indicate that liriodenine is a promising fungicidal compound that should undergo further investigation with some chemical modifications.

Abstract: Background and Aims: The phytochemistry study of Annonaceae has intensified in the last decades due to the discovery of secondary metabolites with antimicrobial activities. The inhibitory activities of extracts and compounds from these species for phytopathogens are less known. The antifungal activity of Sapranthus microcarpus was determined in vitro against six fungal pathogens of important crops. Methods: Hexane, methanol, and alkaloid extracts of the leaves and root and stem bark of S. microcarpus were evaluated to determine whether they could inhibit the mycelial growth of the phytopathogens. The combined activity of the most active extracts for each phytopathogen was also determined. The alkaloid liriodenine was isolated and identified as an antifungal principle. Key results: All extracts inhibited the growth of the six phytopathogens to some extent. The root bark alkaloid extract showed the highest activity. Fusarium oxysporum f. sp. lycopersici and Curvularia lunata were the most sensitive phytopathogens. Liriodenine was active against all the plant pathogens (minimum inhibitory concentrations of 125-500 nmol ml-1). Conclusions: Sapranthus microcarpus is a native natural resource with great phytochemical potential.

Resumen: Antecedentes y Objetivos: El estudio de la fitoquímica de Annonaceae se ha intensificado en las últimas décadas debido al descubrimiento de metabolitos secundarios con actividades antimicrobianas. Las actividades inhibidoras de extractos y compuestos de estas especies para fitopatógenos son menos conocidas. Se determinó la actividad antifúngica in vitro de Sapranthus microcarpus contra seis hongos fitopatógenos de cultivos importantes. Métodos: Se evaluaron extractos metanólicos, hexánicos y alcaloidales de las hojas, y corteza de tallos y raíces de S. microcarpus por el método de inhibición micelial. También se determinó la actividad combinada de los extractos más potentes contra cada fitopatógeno. Se aisló e identificó al alcaloide liriodenina como uno de los metabolitos antifúngicos. Resultados clave: Todos los extractos, en alguna medida, inhibieron el crecimiento de los seis fitopatógenos. Los extractos alcaloidales mostraron la mayor actividad. Fusarium oxysporum f. sp. lycopersici y Curvularia lunata fueron los fitopatógenos más sensibles. Liriodenina fue activo contra todos los fitopatógenos (a concentraciones inhibitorias mínimas de 125-500 nmol ml-1). Conclusiones: Sapranthus microcarpus es un recurso nativo con alto potencial fitoquímico.

Abstract: Background. Phenological descriptions of plant species are used to understand cycles of foliation, flowering and fruiting. These studies provide useful means for understanding the relationships between plants and their environment. Question. What are the the foliar and reproductive strategies of four species of Annona living in the Deciduous Forest of Chiapas, Mexico? Species studied. Annona lutescens, A. macroprophyllata, A. reticulata and A. purpurea. Study sites. Three areas of deciduous forest were studied (16°31’36’’ N, 92°58’25’’ W, 16°28’82’’ N, 92°57’48’’ W, and 16°21’47’’ N, 92°58’32’’ W), located between the towns of Parral and Chiapa de Corzo, Chiapas, Mexico. Method. Stages of foliation, flowering, fruiting and senescence of 20 individuals per species were documented by written and photographic records during the course of a year. Variable time periods were adapted to the extended BBCH scale. The phenology was correlated with the stages of rain and drought by Spearman’s correlations Results. Phenological characteristics distinguish A. reticulata and A. lutescens as species with a reproductive strategy (flowering and fruiting) synchronized with the dry months, in contrast to A. purpurea and A. macroprophyllata, which show a reproductive strategy associated with the rainy season. The foliation of the four species is phenologically related to the rainy season. Conclusions. The species studied could be grouped according to their leaf pattern as brevideciduous or deciduous and according to their reproductive pattern as having long-term or short-term strategies.

Resumen Antecedentes. Las descripciones fenológicas de las especies vegetales son usadas para conocer los ciclos de foliación, floración y fructificación de las plantas cultivadas, estos estudios proveen herramientas útiles para comprender las relaciones entre las plantas y su ambiente. Preguntas. ¿Cuáles serían las estrategias foliares y reproductivas de cuatro especies del género Annona que habitan en la Selva Baja Caducifolia de Chiapas, México? Especies en estudio. Annona lutescens, A. macroprophyllata, A. reticulata y A. purpurea Sitio de estudio. Se estudiaron tres áreas de la selva baja caducifolia, localizada entre los municipios de Chiapas de Corzo y Parral, en Chiapas, México (16° 31’ 36’’ N, 92° 58’ 25’’ W; 16° 28’ 82’ ‘N, 92° 57’ 48’’ W, and 16° 21’ 47’’ N, 92° 58’ 32’’ W). Métodos. Las etapas de foliación, floración, fructificación y senescencia de 20 individuos por especie se documentaron mediante registros fotográficos y escritos durante un año y en periodos de tiempo variable y se adecuaron a la escala BBCH extendida. La fenología y las etapas de lluvia y sequía se asociaron a través de correlaciones de Spearman. Resultados. Las características fenológicas distinguen a A. reticulata y A. lutescens como especies con estrategia de reproducción (floración y fructificación) sincronizadas con los meses de secas, contrastando con las de A. purpurea y A. macroprophyllata que están asociadas a la época de lluvias. La fenología de foliación de las cuatro especies está relacionada con la época de lluvias. Conclusiones. Las especies estudiadas pudieran agruparse de acuerdo a su patrón foliar en brevideciduas y deciduas y de acuerdo a su patrón reproductivo en especies con estrategias largas y cortas

RESUMO Sementes de Annonaceae são conhecidas por possuírem mecanismos de dormência que variam de impermeabilidade de tegumento até à dormência fisiológica. Desta forma, o objetivo deste estudo foi avaliar os efeitos da giberelina GA3 e de GA4+7+ a citocinina benziladenina (GA+BA) na superação da dormência de sementes de Annona macroprophyllata Donn. Sm (papausa) e Annona purpurea Moc. & Sessé ex Dunal (chincuya). O experimento foi realizado com a aplicação de GA3 e GA4+7 + BA nas concentrações de 0; 200, 400, 500, 600, 800 e 1.000 mg L-1. O uso dos reguladores resultou na quebra da dormência em ambas as espécies. Contudo, a aplicação da mistura GA4+7 + BA resultou em maiores incrementos na germinação de A. macroprophyllata do que em A. purpurea. Os tratamentos que promoveram as maiores porcentagens de germinação foram 200 mg L-1 de GA4+7 + BA para A. macroprophyllata (77%) e 200 mg L-1 de GA4+7 + BA e 500 mg L-1 de GA3 para A. purpurea (30% e 29%, respectivamente). O índice de velocidade, o tempo médio e a frequência de germinação foram diferentes para ambas as espécies e com ambos os reguladores. Conclui-se que, apesar de GA3 e GA4+7 + BA promoverem a germinação, a mistura GA4+7 + BA foi mais efetiva que GA3 para quebrar a dormencia de ambas as espécies. Além disso, sementes de A. purpurea apresentam dormência mais difícil de ser superada que A. macroprophyllata.

ABSTRACT Some Annonaceae seeds are known to exhibit dormancy mechanisms ranging from possible seed coat impermeability to physiological dormancy. Thus, the aim of this study was to evaluate the effects of gibberellin (GA) GA3 and GA4+7 + benzyladenine (GA4+7 + BA) application in seeds of Annona macroprophyllata Donn. Sm (papausa) and Annona purpurea Moc. & Sessé ex Dunal (chincuya). The experiment was performed by the application of GA3 and GA4+7 + BA on seeds in concentrations of 0, 200, 400, 500, 600, 800 and 1000 mg L-1. The regulators broke the dormancy of both species. However, application of the GA4+7 + BA mixture had more significant results, with greater increases in germination in A. macroprophyllata than in A. purpurea. Treatments that promoted the highest germinations were GA4+7 + BA at a concentration of 200 mg L-1 for A. macroprophyllata (77%) and 200 mg L-1 of GA4+7 + BA and 500 mg L-1 of GA3 for A. purpurea (30% and 29%, respectively). Rate index, mean time and frequency of germination were distinct for both species and both treatments. Although both GA3 and GA4+7 + BA promote germination, the GA4+7 + BA mixture was more effective than GA3 to overcoming seed dormancy of both species, A. purpurea has a harder dormancy than A. macroprophyllata

Annona macroprophyllata (papausa or ilama) is an economically important species of fruit that is consumed in large quantities in Central America and is considered to have the finest flavour of all of the custard apples (Annonaceae). This species presents propagation problems including the six-month dormancy of the seeds, which has not been fully explained. Among the factors affecting the germination of Annona seeds are the position inside of the fruit and the presence of a micropylar woody plug. The present work addresses the importance of these factors in the germination and dormancy of this species, using a randomised design with seeds from 169 fruits of the white variety of Annona macroprophyllata. The fruits were divided into three sections (basal, middle, and apical) in which seeds were evaluated for nine months with the following parameters: the size and weight of the seeds and the viability and germination percentage considering the presence or absence of the micropylar plug. The results show no correlation between the breaking of dormancy and either the position of the seeds in the fruit or the micropylar plug, but they do establish the importance of the micropylar plug for germination. The breaking of dormancy requires storage for the length of the dry season in the tropical deciduous forest, the habitat of these plants.

Annona macroprophyllata conocida comúnmente como papausa o ilama es una especie frutícola de importancia económica, consumida en el sur de México y en América central; es considerada por su sabor, el fruto más fino de las anonáceas. Esta especie presenta problemas de propagación debido al periodo de latencia de hasta seis meses que tienen sus semillas, el cual no ha sido completamente explicado. Entre los factores que impactan la germinación de las semillas, están la posición de estas dentro del fruto y la presencia de un tapón micropilar leñoso. El presente trabajo investigó la importancia de estos factores relacionados a la germinación y latencia de esta especie. Para este estudio, se estructuró un diseño completamente al azar usando 169 frutos de Annona macroprophyllata de la variedad blanca. Los frutos fueron divididos en tres secciones transversalmente (basal, medio y apical), las semillas de cada sección fueron caracterizadas en tamaño y peso y por nueve meses fueron almacenadas y evaluadas con relación a su viabilidad y capacidad germinativa, considerando al mismo tiempo la presencia o ausencia del tapón micropilar. Los resultados no correlacionan la posición de las semillas en el fruto, ni el tapón micropilar con el rompimiento de la latencia, pero revelan la importancia del tapón micropilar para la germinación. El rompimiento de la latencia de esta especie depende del tiempo de almacenamiento, el cual está completamente asociado a la temporada de sequía de la selva baja caducifolia, donde esta especie habita.

Estudos químicos em plantas da família Annonaceae se intensificaram nas últimas décadas, devido à descoberta de moléculas de anonáceas com potencial medicinal (por exemplo, alcalóides benzilisoquinolina e acetogeninas). Aproximadamente 500 alcaloides têm sido identificados em 138 espécies de Annonaceae em 43 gêneros. Além disso, até 2004, 593 acetogeninas de annonaceas (ACGs) foram identificados, a partir de 51 espécies em 13 gêneros. Isto sugere que as plantas desta família atribuem recursos importantes para a biossíntese destes compostos. Apesar da diversidade destas moléculas, suas funções biológicas, incluindo as suas funções fisiológicas e/ou ecológicas, não são bem compreendidas. Neste estudo, foram fornecidos novos dados que descrevem a variedade e a distribuição de certos alcaloides e ACGs em annonaceas em fases distintas de desenvolvimento. As possíveis relações entre alguns destes compostos e as alterações climáticas sazonais ocorrendo no habitat são também discutidas. Estes dados vão melhorar a nossa compreensão do metabolismo secundário dessas moléculas farmacologicamente importantes e seus padrões de expressão durante o desenvolvimento, o que ajudará a determinar as condições ideais de crescimento e épocas de colheita para a sua produção.

Chemical studies of the plant family Annonaceae have intensified in the last several decades due to the discovery of annonaceous molecules with medicinal potential (e.g., benzylisoquinoline alkaloids and acetogenins). Approximately 500 alkaloids have been identified in 138 Annonaceae species in 43 genera. In addition, until 2004, 593 annonaceous acetogenins (ACGs) had been identified, from 51 species in 13 genera.This suggests that plants from this family allocate important resources to the biosynthesis of these compounds. Despite the diversity of these molecules, their biological roles, including their physiological and/or ecological functions, are not well understood. In this study, it was provided new data describing the variety and distribution of certain alkaloids and ACGs in annonaceous plants in distinct stages of development. The potential relationships among some of these compounds and the seasonally climatic changes occurring in the plant habitat are also discussed. These data will improve our understanding of the secondary metabolism of these pharmacologically important molecules and their expression patterns during development, which will help to determine the optimal growth conditions and harvest times for their production.

Los alcaloides bencilisoquinolínicos (ABI) son metabolitos especializados con una distribución filogenética antigua pero conservada todavía en clados modernos. Varios de ellos, como la morfina, sanguinerina y berberina tienen importancia en la medicina moderna. En esta revisión se analizan los aspectos más sobresalientes del estado actual de la biosíntesis de ABI. Se han realizado estudios que han permitido conocer la biosíntesis de 22 de estos metabolitos nitrogenados. En su formación participan 43 enzimas agrupadas en oxido-reductasas, transferasas y liasas, que en algunos casos representan ejemplos atípicos de la forma en la que se originó la diversificación del metabolismo secundario, entre ellos proteínas citocromo P450 (CYP450) con actividades catalíticas para la ruta de los ABI, o la enzima norcoclaurina sintasa (NCS) que esta emparentada con proteínas alergénicas de defensa. Así mismo, hay avances genéticos en los que se ha podido caracterizar 30 enzimas, permitiendo conocer procesos de regulación. Otro aspecto interesante es la compartimentación de los sitios de biosíntesis y acumulación de ABI ya que en varios casos están separados espacialmente y en distintas especies o en la misma pueden participar varios tipos de células. Ello ha sugerido el transporte intra e intercelular de los alcaloides, los precursores y de las enzimas, se ha documentado el transporte de berberina entre el citoplasma y las vacuolas del almacenamiento. El panorama de la biosíntesis de ABI se ha construido con los estudios de ejemplares de importancia farmacológica.

The benzylisoquinoline alkaloids (BIA) are specialized metabolites with an ancient phylogenetic distribution, but still preserved in modern clades. Some of them, such as morphine, sanguinerine or berberine, are important for modern medicine. This review discusses the highlights of the current state of the biosynthesis of BIA. There have been studies that show the biosynthesis of 22 of these nitrogenous metabolites. In their formation there are 43 enzymes grouped into oxidoreductases, transferases and lyases, which in some cases represent atypical examples of the manner in which the secondary metabolism diversification was originated. Two of these examples are the cytochrome proteins P450 (P450), with catalytic activities for ABI route, or the norcoclaurine synthase enzyme (NCS), which share substantial identity with defense allergenic proteins. Likewise, there are genetic advances that have produced the characterization of 30 enzymes, allowing knowledge of regulatory processes. Another interesting aspect is the compartmentation of the biosynthesis sites and accumulation of BIA, since in several cases they are spatially separated and in different species, or in the same species several types of cells may be involved. This has suggested intra and intercellular transport of alkaloids, precursors and enzymes, and it has been documented berberine transport between the cytoplasm and the vacuoles of storage. The picture for the biosynthesis of BIA has been constructed with exemplary studies of alkaloids with pharmacological importance.

Os alcalóides benzilisoquinolinas (ABI) são metabólitos especializados com uma distribuição filogenética antiga, mas ainda preservada em clados modernos. Vários deles, como a morfina, sanguinarina e berberina são importantes na medicina moderna. Neste artigo, se analisam os aspectos mais destacados do estado atual da biossíntese de ABI; há estudos que tem permitido conhecer a biossíntese de 22 desses metabólitos nitrogenados. Na sua síntese participam 43 enzimas agrupadas em oxidoreductases, transferases, liases e, em alguns casos, representam exemplos atípicos da forma pela qual se originou a diversificação do metabolismo secundário, incluindo as proteínas do citocromo P450 (CYP450), com atividades catalíticas para a rota dos ABI, ou a enzima norcoclaurina sintase (NCS), que está relacionada com proteínas alergênicas de defesa. Da mesma forma, há avanços genéticos na caracterização de 30 enzimas, permitindo conhecer processos de regulação. Outro aspecto interessante é a compartimentalização dos sítios de biossíntese e acumulação de ABI uma vez que em muitos casos estão separados espacialmente e em diferentes espécies, ou na mesma podem participar vários tipos de células. Isto há sugerido o transporte intra e intercelular de alcalóides, precursores das enzimas; tem sido documentado o transporte de berberina entre o citoplasma e os vacúolos de armazenamento. A perspectiva na biossíntese de ABI foi construída com os estudos de exemplares de importância farmacológica.