Journal of Microbiology

Dong-Min Han

10 Articles

Characterization of NpgA, a 4'-phosphopantetheinyl transferase of Aspergillus nidulans, and evidence of its involvement in fungal growth and formation of conidia and cleistothecia for development: Jung-Mi Kim , Ha-Yeon Song , Hyo-Jin Choi , Kum-Kang So , Dae-Hyuk Kim , Keon-Sang Chae , Dong-Min Han , Kwang-Yeop Jahng; J. Microbiol. 2015;53(1):21-31. Published online January 4, 2015; DOI: https://doi.org/10.1007/s12275-015-4657-8

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The null pigmentation mutant (npgA1) in Aspergillus nidulans
results
in a phenotype with colorless organs, decreased branching growth, delayed of asexual spore development, and aberrant cell wall structure. The npgA gene was isolated from A. nidulans to investigate these pleiomorphic phenomena of npgA1 mutant. Sequencing analysis of the complementing gene indicated that it contained a 4􍿁-phosphopantetheinyl transferase (PPTase) superfamily domain. Enzymatic assay of the PPTase, encoded by the npgA gene, was implemented in vivo and in vitro. Loss-of-function of LYS5, which encoded a PPTase in Saccharomyces cerevisiae, was functionally complemented by NpgA, and Escherichia coli-derived NpgA revealed phosphopantetheinylation activity with the elaboration of 3􍿁5􍿁-ADP. Deletion of the npgA gene caused perfectly a lethal phenotype and the absence of asexual/sexual sporulation and secondary metabolites such as pigments in A. nidulans. However, a cross feeding effect with A. nidulans wild type allowed recovery from deletion defects, and phased-culture filtrate from the wild type were used to verify that the npgA gene was essential for formation of metabolites needed for development as well as growth. In addition, forced expression of npgA promoted the formation of conidia and cleistothecia as well as growth. These results indicate that the npgA gene is involved in the phosphopantetheinylation required for primary biological processes such as growth, asexual/sexual development, and the synthesis of secondary metabolites in A. nidulans.

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Purification-free antifungal biocontrol platform using engineered Saccharomyces cerevisiae secreting Trichoderma atroviride chitinase
Ha-Yeon Song, Dae-Hyuk Kim, Jung-Mi Kim, Ji Young Kang
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Regulators of the Asexual Life Cycle of Aspergillus nidulans
Ye-Eun Son, Jae-Hyuk Yu, Hee-Soo Park
Cells.2023; 12(11): 1544. CrossRef
De novo biosynthesis of carminic acid in Saccharomyces cerevisiae
Qian Zhang, Xinglong Wang, Weizhu Zeng, Sha Xu, Dong Li, Shiqin Yu, Jingwen Zhou
Metabolic Engineering.2023; 76: 50. CrossRef
Expanding luciferase reporter systems for cell-free protein expression
Wakana Sato, Melanie Rasmussen, Christopher Deich, Aaron E. Engelhart, Katarzyna P. Adamala
Scientific Reports.2022;[Epub] CrossRef
Biosynthesis of fungal polyketides by collaborating and trans-acting enzymes
Elizabeth Skellam
Natural Product Reports.2022; 39(4): 754. CrossRef
Liamocins biosynthesis, its regulation in Aureobasidium spp., and their bioactivities
Xin-Xin Kang, Shu-Lei Jia, Xin Wei, Mei Zhang, Guang-Lei Liu, Zhong Hu, Zhe Chi, Zhen-Ming Chi
Critical Reviews in Biotechnology.2022; 42(1): 93. CrossRef
Shimalactone Biosynthesis Involves Spontaneous Double Bicyclo‐Ring Formation with 8π‐6π Electrocyclization
Isao Fujii, Makoto Hashimoto, Kaori Konishi, Akiko Unezawa, Haruka Sakuraba, Kenta Suzuki, Harue Tsushima, Miho Iwasaki, Satsuki Yoshida, Akane Kudo, Rina Fujita, Aika Hichiwa, Koharu Saito, Takashi Asano, Jun Ishikawa, Daigo Wakana, Yukihiro Goda, Ayumi
Angewandte Chemie.2020; 132(22): 8542. CrossRef
Shimalactone Biosynthesis Involves Spontaneous Double Bicyclo‐Ring Formation with 8π‐6π Electrocyclization
Isao Fujii, Makoto Hashimoto, Kaori Konishi, Akiko Unezawa, Haruka Sakuraba, Kenta Suzuki, Harue Tsushima, Miho Iwasaki, Satsuki Yoshida, Akane Kudo, Rina Fujita, Aika Hichiwa, Koharu Saito, Takashi Asano, Jun Ishikawa, Daigo Wakana, Yukihiro Goda, Ayumi
Angewandte Chemie International Edition.2020; 59(22): 8464. CrossRef
Genomic analysis of a riboflavin-overproducing Ashbya gossypii mutant isolated by disparity mutagenesis
Tatsuya Kato, Junya Azegami, Ami Yokomori, Hideo Dohra, Hesham A. El Enshasy, Enoch Y. Park
BMC Genomics.2020;[Epub] CrossRef
Genetic evidences for the core biosynthesis pathway, regulation, transport and secretion of liamocins in yeast-like fungal cells
Si-Jia Xue, Guang-Lei Liu, Zhe Chi, Zhi-Chao Gao, Zhong Hu, Zhen-Ming Chi
Biochemical Journal.2020; 477(5): 887. CrossRef
A Novel Rapid Fungal Promoter Analysis System Using the Phosphopantetheinyl Transferase Gene,npgA, inAspergillus nidulans
Ha-Yeon Song, Dahye Choi, Dong-Min Han, Dae-Hyuk Kim, Jung-Mi Kim
Mycobiology.2018; 46(4): 429. CrossRef
Both a PKS and a PPTase are involved in melanin biosynthesis and regulation of Aureobasidium melanogenum XJ5-1 isolated from the Taklimakan desert
Hong Jiang, Guang-Lei Liu, Zhe Chi, Jian-Ming Wang, Ly-Ly Zhang, Zhen-Ming Chi
Gene.2017; 602: 8. CrossRef
Identification of a Polyketide Synthase Gene in the Synthesis of Phleichrome of the Phytopathogenic Fungus Cladosporium phlei
Kum-Kang So, Yun-Jo Chung, Jung-Mi Kim, Beom-Tae Kim, Seung-Moon Park, Dae-Hyuk Kim
Molecules and Cells.2015; 38(12): 1105. CrossRef

A Putative APSES Transcription Factor Is Necessary for Normal Growth and Development of Aspergillus nidulans: Ji-Yeon Lee , Lee-Han Kim , Ha-Eun Kim , Jae-Sin Park , Kap-Hoon Han , Dong-Min Han; J. Microbiol. 2013;51(6):800-806. Published online December 19, 2013; DOI: https://doi.org/10.1007/s12275-013-3100-2

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Abstract PDF

The nsdD gene encoding a GATA type transcription factor positively controls sexual development in Aspergillus nidulans. According to microarray data, 20 genes that were upregulated by deleting nsdD during various life cycle stages were randomly selected and deleted for functional analysis. None of the mutants showed apparent changes in growth or development compared with those of the wild-type except the AN3154 gene that encodes a putative APSES transcription factor and is an ortholog of Saccharomyces cerevisiae swi4. Deleting AN3154 resulted in retarded growth and development, and the gene was named rgdA (retared growth and development). The rgdA deletion mutant developed a reduced number of conidia even under favorable conditions for asexual development. The retarded growth and development was partially suppressed by the veA1 mutation. The conidial heads of the mutant aborted, showing reduced and irregular shaped phialides. Fruiting body development was delayed compared with that in the wild-type. The mutant did not respond to various nutritional or environmental factors that affected the development patterns. The rgdA gene was expressed at low levels throughout the life cycle and was not significantly affected by several regulators of sexual and asexual development such as nsdD, veA, stuA, or brlA. However, the rgdA gene affected brlA and abaA expression, which function as key regulators of asexual sporulation, suggesting that rgdA functions upstream of those genes.

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AoMbp1 interacts with AoSwi6 and regulates hyphal development, stress tolerance, trap morphogenesis, and pathogenicity in Arthrobotrys oligospora
Na Bai, Hui Yuan, Guiqiu Luo, Yi Chen, Meihua Xie, Jinkui Yang
Pesticide Biochemistry and Physiology.2026; 218: 106930. CrossRef
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Xiao Li, Qianqian Zhang, Qili Liu, Xiaobin Xu, Jinzhu Li, Dandan Zhu, Yuanyuan Zong, Huali Xue, Yang Bi
Postharvest Biology and Technology.2025; 227: 113628. CrossRef
Genome-Wide Mapping Reveals an Extensive AtfA Regulatory Influence on Development, Metabolism, and Stress Preparedness in Aspergillus nidulans
Márton Miskei, Sandugash Ibragimova, Beatrix Kocsis, Tibor Nagy, Hee-Soo Park, Tamás Emri, Jae-Hyuk Yu, Éva Leiter, István Pócsi
Cells.2025; 14(24): 1965. CrossRef
Putative APSES family transcription factor mbp1 plays an essential role in regulating cell wall synthesis in the agaricomycete Pleurotus ostreatus
Hayase Kojima, Moriyuki Kawauchi, Yuitsu Otsuka, Kim Schiphof, Kenya Tsuji, Akira Yoshimi, Chihiro Tanaka, Shigekazu Yano, Takehito Nakazawa, Yoichi Honda
Fungal Genetics and Biology.2024; 175: 103936. CrossRef
Regulators of the Asexual Life Cycle of Aspergillus nidulans
Ye-Eun Son, Jae-Hyuk Yu, Hee-Soo Park
Cells.2023; 12(11): 1544. CrossRef
Characterization of the mbsA Gene Encoding a Putative APSES Transcription Factor in Aspergillus fumigatus
Yong-Ho Choi, Sang-Cheol Jun, Min-Woo Lee, Jae-Hyuk Yu, Kwang-Soo Shin
International Journal of Molecular Sciences.2021; 22(7): 3777. CrossRef
The Putative APSES Transcription Factor RgdA Governs Growth, Development, Toxigenesis, and Virulence in Aspergillus fumigatus
Sang-Cheol Jun, Yong-Ho Choi, Min-Woo Lee, Jae-Hyuk Yu, Kwang-Soo Shin, Aaron P. Mitchell
mSphere.2020;[Epub] CrossRef
Analogous and Diverse Functions of APSES-Type Transcription Factors in the Morphogenesis of the Entomopathogenic Fungus Metarhizium rileyi
Caiyan Xin, Jinping Zhang, Siji Nian, Guangxi Wang, Zhongkang Wang, Zhangyong Song, Guangwei Ren, Ning-Yi Zhou
Applied and Environmental Microbiology.2020;[Epub] CrossRef
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Chenghua Zhang, Gangzheng Wang, Wangqiu Deng, Taihui Li
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Larissa V G Longo, Stephanie C Ray, Rosana Puccia, Chad A Rappleye
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Guangshan Yao, Feng Zhang, Xinyi Nie, Xiuna Wang, Jun Yuan, Zhenhong Zhuang, Shihua Wang
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Isolation and Characterization of Two Methyltransferase Genes, AfuvipB and AfuvipC in Aspergillus fumigatus
Mohammed A. Abdo Elgabbar, Kap-Hoon Han
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Depletion of ε-COP in the COPI Vesicular Coat Reduces Cleistothecium Production inAspergillus nidulans
Eun-Hye Kang, Eun-Jung Song, Jun Ho Kook, Hwan-Hee Lee, Bo-Ri Jeong, Hee-Moon Park
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Hokyoung Son, Myung-Gu Kim, Suhn-Kee Chae, Yin-Won Lee
Journal of Microbiology.2014; 52(11): 930. CrossRef
Transcriptional regulation of fksA, a β-1,3-glucan synthase gene, by the APSES protein StuA during Aspergillus nidulans development
Bum-Chan Park, Yun-Hee Park, Soohyun Yi, Yu Kyung Choi, Eun-Hye Kang, Hee-Moon Park
Journal of Microbiology.2014; 52(11): 940. CrossRef

Isolation and Characterization of Self-Fertile Suppressors from the Sterile nsdD Deletion Mutant of Aspergillus nidulans: Dong-Beom Lee , Lee Han Kim , Jin-Pyo Kim , Kap-Hoon Han , Dong-Min Han; J. Microbiol. 2011;49(6):1054-1057. Published online December 28, 2011; DOI: https://doi.org/10.1007/s12275-011-1111-4

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Abstract PDF

To identify downstream and/or interactive factors of the nsdD gene, which encodes a positive regulator of sexual development of Aspergillus nidulans, suppressor mutants displaying a self-fertile phenotype were isolated from a sterile nsdD deletion mutant. At least five different loci (sndA-E) were identified and genetically analyzed. In the nsdD+ background, most of the suppressors showed a marked increment of sexual development, even under the stress conditions that normally inhibited sexual development. The common phenotype of the suppressor mutants suggested the involvement of the snd genes in the negative regulation of sexual development in response to the environmental factors.

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Inactivation of the global regulator LaeA in Monascus ruber results in a species-dependent response in sporulation and secondary metabolism
Qingpei Liu, Li Cai, Yanchun Shao, Youxiang Zhou, Mu Li, Xiaohong Wang, Fusheng Chen
Fungal Biology.2016; 120(3): 297. CrossRef
A putative APSES transcription factor is necessary for normal growth and development of Aspergillus nidulans
Ji-Yeon Lee, Lee-Han Kim, Ha-Eun Kim, Jae-Sin Park, Kap-Hoon Han, Dong-Min Han
Journal of Microbiology.2013; 51(6): 800. CrossRef
Isolation and Characterization of the gtfA Gene Encoding GAL4-Like Transcription Factor in Aspergillus nidulans
Jae-Sin Park, Dong-Min Han
The Korean Journal of Microbiology.2013; 49(1): 8. CrossRef

The MpkB MAP Kinase Plays a Role in Post-karyogamy Processes as well as in Hyphal Anastomosis During Sexual Development in Aspergillus nidulans: Sang-Cheol Jun , Sei-Jin Lee , Hyun-Joo Park , Ji-Young Kang , Young-Eun Leem , Tae-Ho Yang , Mi-Hee Chang , Jung-Mi Kim , Seung-Hwan Jang , Hwan-Gyu Kim , Dong-Min Han , Keon-Sang Chae , Kwang-Yeop Jahng; J. Microbiol. 2011;49(3):418-430. Published online June 30, 2011; DOI: https://doi.org/10.1007/s12275-011-0193-3

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Two genes encoding MAP kinase homologs, designated as mpkB and mpkC, were isolated from Aspergillus nidulans by PCR with degenerate primers. Deletion and over-expression mutants of mpkC showed no detectable phenotypes under any external stress tested. Deletion of mpkB caused pleiotropic phenotypes including a failure in forming cleistothecia under any induction conditions for sexual development, increased Hülle cell production, slow hyphal growth and aberrant conidiophore morphology. Over-expression of mpkB led to increased cleistothecium production. While the transcripts of mpkB and mpkC were constitutively synthesized through the entire life cycle, their size and amount differed with developmental stages. An outcross test using fluorescent protein reporters showed that the mpkB deletion mutant could not form heterokaryons with wild type. Protoplast fusion experiments showed that the fusant of the mpkB mutant with wild type could undergo normal sexual development. However, heterokaryotic mycelia that were produced from a fusant between two mpkB deletion mutants could not form cleistothecia, although they did appear to form diploid nuclei. These results suggest that the MpkB MAP kinase is required for some post-karyogamy process as well as at the hyphal anastomosis stage to accomplish sexual development successfully.

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Characterization of NpgA, a 4′-phosphopantetheinyl transferase of Aspergillus nidulans, and evidence of its involvement in fungal growth and formation of conidia and cleistothecia for development
Jung-Mi Kim, Ha-Yeon Song, Hyo-Jin Choi, Kum-Kang So, Dae-Hyuk Kim, Keon-Sang Chae, Dong-Min Han, Kwang-Yeop Jahng
Journal of Microbiology.2015; 53(1): 21. CrossRef
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Simple Identification of veA1 Mutation in Aspergillus nidulans: Kap-Hoon Han , Jae-Sin Park , Keon Sang Chae , Dong-Min Han; J. Microbiol. 2010;48(6):885-887. Published online January 9, 2011; DOI: https://doi.org/10.1007/s12275-010-0506-y

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Abstract PDF

The veA gene plays an important role in development of a homothallic filamentous fungus Aspergillus nidulans. The veA1 phenotype can be difficult to distinguish from the wild-type veA. Despite the importance of the veA allele, no efficient identification method has been reported besides DNA sequencing. Here, we present simple physiological and molecular biological ways to distinguish between the veA wild-type and veA1 allele. The novel approaches, which involve incubation in the presence of oxalic acid, polymerase chain reaction using double mismatched primers, and BstXI enzyme digestion, are simpler, faster and more cost-efficient than genome sequencing.

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Chemical composition of essential oil and antifungal activity of Artemisia persica Boiss. from Iran
Reza Dehghani Bidgoli
Journal of Food Science and Technology.2021; 58(4): 1313. CrossRef
Survival factor SvfA plays multiple roles in differentiation and is essential for completion of sexual development in Aspergillus nidulans
Joo-Yeon Lim, Eun-Hye Kang, Yun-Hee Park, Jun-Ho Kook, Hee-Moon Park
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HMGB proteins are required for sexual development in Aspergillus nidulans
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NapA Mediates a Redox Regulation of the Antioxidant Response, Carbon Utilization and Development in Aspergillus nidulans
Ariann E. Mendoza-Martínez, Fernando Lara-Rojas, Olivia Sánchez, Jesús Aguirre
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Zoltán Németh, Ákos Molnár, Balázs Fejes, Levente Novák, Levente Karaffa, Nancy Keller, Erzsébet Fekete
Toxins.2016; 8(12): 354. CrossRef
The SrkA Kinase Is Part of the SakA Mitogen-Activated Protein Kinase Interactome and Regulates Stress Responses and Development in Aspergillus nidulans
Rafael Jaimes-Arroyo, Fernando Lara-Rojas, Özgür Bayram, Oliver Valerius, Gerhard H. Braus, Jesús Aguirre
Eukaryotic Cell.2015; 14(5): 495. CrossRef
A putative APSES transcription factor is necessary for normal growth and development of Aspergillus nidulans
Ji-Yeon Lee, Lee-Han Kim, Ha-Eun Kim, Jae-Sin Park, Kap-Hoon Han, Dong-Min Han
Journal of Microbiology.2013; 51(6): 800. CrossRef
FlbD, a Myb Transcription Factor of Aspergillus nidulans, Is Uniquely Involved in both Asexual and Sexual Differentiation
Jenny Arratia-Quijada, Olivia Sánchez, Claudio Scazzocchio, Jesús Aguirre
Eukaryotic Cell.2012; 11(9): 1132. CrossRef

Predicting the Chemical Composition and Structure of Aspergillus nidulans Hyphal Wall Surface by Atomic Force Microscopy: Hyun-uk Lee , Jong Bae Park , Haeseong Lee , Keon-Sang Chae , Dong-Min Han , Kwang-Yeop Jahng; J. Microbiol. 2010;48(2):243-248. Published online May 1, 2010; DOI: https://doi.org/10.1007/s12275-010-8094-4

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Abstract PDF

In fungi, cell wall plays an important role in growth and development. Major macromolecular constituents of the aspergilli cell wall are glucan, chitin, and protein. We examined the chemical composition and structure of the Aspergillus nidulans hyphal wall surface by an atomic force microscope (AFM). To determine the composition of the cell wall surface, the adhesion forces of commercially available β-glucan, chitin, and various proteins were compared to those of corresponding fractions prepared from the hyphal wall. In both setups, the adhesion forces of β-glucan, chitin, and protein were 25-50, 1000-3000, and 125-300 nN, respectively. Adhesion force analysis demonstrated that the cell surface of the apical tip region might contain primarily chitin and β-glucan and relatively a little protein. This analysis also showed the chemical composition of the hyphal surface of the mid-region would be different from that of the apical region. Morphological images obtained by the tapping mode of AFM revealed that the hyphal tip surface has moderate roughness.

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Screening of Growth- or Development-related Genes by Using Genomic Library with Inducible Promoter in Aspergillus nidulans: Bang-Yong Lee , Sang-Yong Han , Han Gil Choi , Jee Hyun Kim , Kap-Hoon Han , Dong-Min Han; J. Microbiol. 2005;43(6):523-528.; DOI: https://doi.org/2295 [pii]

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Abstract PDF: Using the genomic library constructed at the downstream of the niiA promoter, which induces the over-expression of an inserted DNA fragment, we have attempted to screen the genes affecting growth or development by over-expression. The wild-type strain was transformed using the AMA-niiA(p) library and cultured on 1.2 M sorbitol media, in which asexual sporulation is induced, but sexual development is repressed. Over 100,000 strains transformed to pyrG+ were analyzed with regard to any changes in phenotype. Consequently, seven strains were isolated for further analyses. These strains were designated NOT [niiA(p) over-expression transformants] stains. Four of the strains were of the inducible type, and the remaining strains were of the multi-copy suppression type. Two of the inducible-type strains, NOT1 and NOT40, harbored genes which had been inserted in reverse direction, suggesting that the mutant phenotypes had been derived from an excess amount of anti-sense mRNA. Domain analyses of the deduced polypeptides from the DNA fragments rescued from the transformants revealed that NOT1, NOT40 and NOT6 harbored a LisH motif, a forkhead domain, and a Zn(II)2Cys6 binuclear zinc cluster, respectively.

Regulation of nsdD Expression in Aspergillus nidulans: Kap Hoon Han , Kyu-Yong Han , Min-Su Kim , Dong-Beom Lee , Jong-Hak Kim , Suhn-Kee Chae , Keon-Sang Chae , Dong-Min Han; J. Microbiol. 2003;41(3):259-261.

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Abstract PDF: The nsdD gene has been predicted to encode a GATA type transcription factor with the type IVb zinc finger DNA binding domain functions in activating sexual development of A. nidulans. In several allelic mutants of nsdD producing truncated NsdD polypeptides lacking the C-terminal zinc finger, the transcription level of nsdD gene was greatly increased. Also in an over-expressed mutant, the transcription under its own promoter was reduced. These results suggest that the expression of nsdD is negatively autoregulated. When the nsdD gene was over-expressed, cleistothecia were formed in excess amounts even in the presence of 0.6M KCl that inhibited sexual development of the wild type. Northern blot analysis revealed that the expression of nsdD was repressed by 0.6M KCl. These results strongly suggest that the inhibition of sexual development by salts was carried out via the nsdD involved regulatory network.

Ultrastructure of the Cell Wall of a Null Pigmentation Mutant, npgA1, in Aspergillus nidulans: Yun Shin Chung , § , Jung-Mi Kim , Dong-Min Han , Keon-Sang Chae , Kwang-Yeop Jahng; J. Microbiol. 2003;41(3):224-231.

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Abstract PDF: The null pigmentation mutant (npgA1) of Aspergillus nidulans was previously characterized by its production of no pigment at any stage of its life cycle, its reduction in hyphal branching, and its delay in the asexual spore development. The chemical composition of the cell wall was also altered in npgA1 mutants that became more sensitive to Novozyme 234_TM, which is possibly due to a structural defect in the cell wall. To investigate the effects of the cell wall structure on these pleiomorphic phenomena, we examined the ultrastructure of the cell wall in the npgA1 mutant (WX17). Scanning electron micrographs (SEM) showed that after being cultured for six days, the outermost layer of the conidial wall of WX17 peeled off. Although this phenotype suggested that the cell wall structure in WX17 may be modified, examination using TEM of the fine structure of cross-sectioned hyphal wall of WX17 did not show any differences from that of FGSC4. However, staining for carbohydrates of wall layers showed that the electron-translucent layer of the cell wall was missing in WX17. In addition, the outermost layer H1 of the hyphal wall was also absent in WX17. The ultrastructural observation and cytochemical analysis of cell walls suggested that the pigmentation defect in WX17 may be attributed to the lack of a layer in the cell wall.

Promotion of Asexual Development and Inhibition of Sexual Development of Aspergillus nidulans by Short-Chain Primary Amines: Myung Hoon Song , Kuppusamy Selvam , Chang-Jun Choi , Kwang-Yeop Jahng , Dong-Min Han , Keon-Sang Chae; J. Microbiol. 2002;40(3):230-233.

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Abstract PDF: Effects of short-chain primary amines on Aspergillus nidulans development were analyzed. Propylamine induced asexual development and inhibited sexual development. Even on medium containing lactose as the sole carbon source, on which little conidial heads are formed and sexual structures are formed preferentially, or when sexual development was induced, propylamine induced asexual development and inhibited sexual development. These effects of propylamine seemed to be due to accumulation of mRNA of the brlA gene, which has been identified as a positive regulator of asexual development, and due to the reduction of the veA mRNA level. The veA gene has been identified as an activator of sexual development and also as an inhibitor of asexual development. Other primary amines, methylamine and ethylamine, showed identical effects on development where short-chain primary amine also promoted asexual development and inhibited sexual development.

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