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Interaksi Allelopathic dan Allelochemicals: Kemungkinan Baru untuk Berkelanjutan Manajemen Gulma

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Interaksi Allelopathic dan Allelochemicals: Baru
Kemungkinan Pengelolaan Gulma Berkelanjutan
H. P. Singh a, Daizy R. Batish a & R.K. Kohli a b
Departemen Botani, Universitas Punjab, Chandigarh 160 014, India
b Pusat Lingkungan dan Studi Vokasional, Universitas Punjab, Chandigarh, India
Diterbitkan online: 18 Jun 2010.

To cite this article: H. P. Singh , Daizy R. Batish & R.K. Kohli (2003) Allelopathic Interactions and Allelochemicals: New
Possibilities for Sustainable Weed Management, Critical Reviews in Plant Sciences, 22:3-4, 239-311, DOI: 10.1080/713610858

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 Critical Reviews in Plant Sciences, 22(3&4):239–311 (2003)

Allelopathic Interactions and Allelochemicals: New

Possibilities for Sustainable Weed Management
H. P. Singh,1 Daizy R. Batish,1 and R.K. Kohli1,2*
1
Department of Botany, Panjab University, Chandigarh 160 014, India; 2Centre for Environment and Vocational
Studies, Panjab University, Chandigarh, India

Referee: Dr. Amrjits S. Basra, Central Plains Crop Technology, 5912 North Meridian Avenue, Wichita, KS 67204

* Corresponding author (Email: rkkohli45@yahoo.com)

Downloaded by [Central University of Punjab] at 04:35 07 January 2015

TABLE OF CONTENTS

I. Weeds – Characteristics, Adaptive Strategies and Roles in

Agroecosystems ...................................................................................... 240

II. Searching the Potential of Allelopathic Interactions in

Agroecosystems ...................................................................................... 242

III. Cropping Patterns, Allelopathy and Weed Management ................. 244

A. Crop Rotation .................................................................................... 244
B. Cover and Smother crops ................................................................. 245
1. Rye (Secale cereale L.) as a Model Cover Crop .......................248
C. Green Manure Crops .........................................................................249
D. Crop Residues — Wheat, Rice, Sorghum, Alfalfa, and Other
Crop Residues ......................................................................................... 250

IV. Screening of Crop Varieties for Allelopathic Strength .....................

253

V. Allelochemicals — Nature’s Own Herbicides ....................................

261
A. From Higher Plants (Terpenoids, Benzoxazinoids,
Glucosinolates, Quassinoids, Cyanogenic Glycosides, Saponins,
Sorgoleone, Juglone
and Other compounds) ............................................................................ 262
B. From Microbes .................................................................................. 271
1. Microbial Phytotoxins ................................................................ 272
2. Direct Use of Microbes for Weed Management .......................282
239
 0735-2689/03/$.50
© 2003 by CRC Press LLC

240
 VI. Potential of Allelopathy for Parasitic Weed Management ............... 283
1. Potential of Allelopathy, Through Allelochemicals and Cultural
Practices ...................................................................................... 283
2. Control of Parasitic Weeds Through Microbes ......................... 285

VII. Management of Aquatic Weeds Through Allelopathy ...................... 286

VIII. Role of Molecular Genetic Techniques for Improving Crops for

Allelopathic Traits .................................................................................
287

References ......................................................................................................... 289

ABSTRAK: Gulma diketahui menyebabkan kerugian besar karena gangguan mereka di

agroekosistem. Karena masalah lingkungan dan kesehatan manusia, upaya dunia sedang
dilakukan untuk mengurangi ketergantungan berat pada herbisida sintetis yang digunakan
untuk mengendalikan gulma. Dalam hal ini fenomena alelopati, yang diekspresikan melalui
pelepasan bahan kimia oleh tanaman, telah disarankan untuk menjadi salah satu alternatif
yang mungkin untuk mencapai manajemen gulma berkelanjutan. Penggunaan alelopati
untuk mengendalikan gulma dapat berupa secara langsung memanfaatkan interaksi
alelopati alami, terutama tanaman tanaman, atau dengan menggunakan allelochemicals
sebagai herbisida alami. Dalam kasus sebelumnya, sejumlah tanaman tanaman dengan
potensi allelopathic dapat digunakan sebagai penutup, smother, dan tanaman pupuk hijau
untuk mengelola gulma dengan membuat manipulasi yang diinginkan dalam praktik budaya
dan pola tanam. Ini dapat dirotasi atau ditumpangsusun dengan tanaman utama untuk
mengelola gulma sasaran (termasuk yang parasit) secara selektif. Bahkan mulsa / residu
tanaman juga dapat memberikan manfaat yang diinginkan. Tidak hanya gulma darat,
bahkan alelopati dapat dimanipulasi dengan tepat untuk pengelolaan gulma air. The
allelochemicals hadir di tanaman yang lebih tinggi serta di mikroba dapat langsung
digunakan untuk manajemen gulma pada pola herbisida. Bioefficacy mereka dapat
ditingkatkan dengan perubahan struktural atau sintesis analog kimia berdasarkan pada
mereka. Selanjutnya, untuk meningkatkan potensi tanaman allelopathic, beberapa
perbaikan dapat dilakukan dengan penggunaan bioteknologi atau genomik dan proteomik.
Dalam konteks ini baik produksi alelokimia dapat ditingkatkan atau transgenik dengan
pengkodean gen asing untuk alelokimia penekan gulma tertentu dapat diproduksi. Pada
yang pertama, pemuliaan konvensional dan teknik genetika molekuler berguna. Namun,
dengan pemuliaan konvensional yang lambat dan sulit, lebih ditekankan pada penggunaan
teknik modern seperti penanda molekuler dan pemilihan dibantu oleh mereka. Meskipun
kemajuan dalam hal ini lambat, namun beberapa hasil yang menjanjikan akan datang dan
lebih banyak diharapkan di masa depan. Ulasan ini mencoba untuk membahas semua
aspek alelopati ini untuk pengelolaan gulma yang berkelanjutan.

KATA KUNCI: agroekosistem, alelokimia, alelopati, sarana bioteknologi, tanaman penutup, perbaikan tanaman, sisa tanaman, pola tanam,
tanaman pupuk hijau, herbisida alami, tanaman cabul, pertanian berkelanjutan, manajemen gulma.

241
 I. WEEDS — CHARACTERISTICS,
ADAPTIVE STRATEGIES, AND ROLES
IN AGROECOSYSTEMS
Kata gulma berarti tumbuhan liar
yang tumbuh di tempat yang tidak
diinginkan terutama di ladang
atau di kebun yang mengganggu
pertumbuhan tanaman yang
dibudidayakan. Sejumlah definisi
gulma telah diusulkan, tetapi
tidak ada yang mencapai
kepuasan universal. Gulma dapat
didefinisikan sebagai tanaman
Downloaded by [Central University of Punjab] at 04:35 07 January 2015
242
dengan nilai ekonomi kecil dan memiliki potensi
untuk menjajah habitat yang terganggu atau yang
dimodifikasi oleh aktivitas manusia. Aldrich
(1984) mendefinisikan gulma sebagai tanaman
yang berasal dari lingkungan alami dan sebagai
respons terhadap manusia (dikenakan) atau
kondisi alam mengganggu tanaman dan aktivitas
manusia. Gulma juga telah didefinisikan sebagai
tanaman yang tidak menyenangkan yang
mengganggu kegiatan dan kesejahteraan manusia
(WSSA,
1994). Cousens and Mortimer (1995) dijelaskan
 gulma sebagai tanaman vaskular yang merupakan
hambatan alami untuk aktivitas manusia atau
kesehatan atau yang diketahui menyebabkan
perubahan yang tidak dapat diterima untuk komunitas
tumbuhan alami. Mohler (2001) meruntuhkan gulma
sebagai tanaman yang secara khusus berhasil dalam
penjajahan yang terganggu (tetapi berpotensi
produktif) dan mempertahankan kelimpahannya di
bawah kondisi terganggu yang berulang. Sebagian
besar definisi tentang gulma memiliki satu kesamaan
bahwa mereka adalah tamu yang tidak diinginkan di
area budidaya dan perlu dikontrol atau dikelola. Secara
ekologis, gulma merupakan pelopor dari suksesi
sekunder, sedangkan dari perspektif ekonomi mereka
adalah tanaman yang kebajikannya belum dinilai
sepenuhnya. Gulma juga telah dianggap sebagai
sumber potensial keragaman genetik dan kerabat liar
tanaman tanaman, dan karenanya harus dipertahankan
untuk program perbaikan tanaman di masa mendatang
(Hammer et al., 1997).
Gulma hanya mewakili 0,1% dari flora dunia
dan dengan demikian tidak merupakan bagian terbesar.
Di agroecosystems gulma dan tanaman telah berevolusi
bersama-sama langsung dari zaman prasejarah seperti
yang diungkapkan oleh studi analisis serbuk sari (Cousens
dan Mortimer, 1995). Pembiayaan mereka juga dibuktikan
dari fakta bahwa lima keluarga besar (Poaceae,
Solanaceae, Euphorbiaceae, Convolvulaceae, dan
Fabaceae) di mana gulma lama juga menyediakan 75%
pasokan makanan dunia (Holm et al., 1977). Hammer
(1988) telah menunjukkan bahwa meskipun baik gulma
dan tanaman telah berevolusi di bawah garis evolusi yang
serupa, tetapi manusia menciptakan lingkungan yang
menguntungkan hanya untuk tanaman / tanaman yang
berguna. Gulma memiliki banyak asal; beberapa
diperkenalkan dari wilayah geografis lain untuk beberapa
tujuan yang berguna dan kemudian menganggap proporsi
yang kurus, sedangkan yang lain menginvasi secara tidak
sengaja. Sebagai contoh, di India Lantana camara L.
diperkenalkan sebagai semak hias, tetapi sekarang
diasumsikan memiliki posisi gulma yang serius. Di sisi
lain, ragweed Parthenium (Parthenium hysterophorus L.)
telah memasuki India secara tiba-tiba dan sekarang
merupakan salah satu gulma berbahaya. Tidak hanya
eksotik tetapi bahkan spesies asli menjadi kurus karena
gangguan manusia yang berlebihan.
Gulma menyebabkan sejumlah bahaya di agro-
ekosistem. Karena gangguan mereka dengan
tanaman (baik yang kompetitif dan alelopati),
mereka mengurangi hasil panen yang
menyebabkan kerugian besar dalam skala global.
Sekitar 240 spesies gulma dilaporkan bersifat
allelopathic dan mengganggu pertumbuhan dan
produksi tanaman (Qasem dan Foy, 2001).
Selain itu, mereka merusak kualitas tanaman,
menyumbat saluran air, menyebabkan masalah
kesehatan pada manusia, dan terlihat tidak sedap
dipandang di area yang menyenangkan seperti
taman, taman, jalur, dan trotoar, dll. Selanjutnya,
biaya eradikasi gulma juga sangat besar.
Piementel et al. (2001) telah menunjukkan
bahwa di gulma AS menyebabkan sekitar 12%
kerugian dalam hasil panen dan biayanya hampir
US $ 35 miliar untuk mengendalikannya.
Biayanya bahkan lebih banyak di negara
berkembang. Selain merusak hasil panen dan
tidak menarik, mereka juga menyebabkan
bahaya kebakaran (Zimdahl, 1999). Gulma juga
dapat menjangkiti serangga dan patogen,
menambahkan lebih banyak komplikasi ke
kontrol mereka.
Mengingat karakteristik ini dari gulma dan
bahaya mereka, menjadi sangat penting untuk
mengendalikan mereka. Sejumlah praktik
manajemen tersedia. Di masa sebelumnya,
karena tidak ada bahan kimia sintetis yang
diketahui, rotasi tanaman, polikultur, dan praktik
manajemen lainnya diadili dengan input rendah
tetapi berkelanjutan. Dengan penemuan herbisida
sintetis pada awal 1930-an, ada pergeseran dalam
praktik manajemen gulma menuju input tinggi
dan berorientasi pada target. DNOC (4,6-dinitro-
o-cresol) adalah herbisida sintetik pertama yang
dipatenkan, dan kemudian sejumlah pengatur
pertumbuhan sintetis seperti 2,4-D (2,4-
dichlorophenoxyacetic acid) dan MCPA (2-
methyl-4-chlorophenoxyacetic acid) secara
keseluruhan menggantikan herbisida lain karena
sifatnya yang selektif dan dengan demikian
menjadi dasar industri agrokimia. Ini telah
meningkatkan produksi tanaman dengan
meminimalkan persaingan antara tanaman dan
gulma dan mengurangi risiko gangguan dari
gulma (Parry, 1989). Karena itu, praktik
pertanian mulai sangat bergantung pada bahan
kimia ini. Di AS saja, 75% perlindungan
tanaman didasarkan pada input herbisida (Duke,
1999). Namun, selama tahun 1980-an kontrol
gulma kimia mulai menjadi tidak populer di
seluruh dunia karena meningkatnya biaya dan
berbagai bahaya lingkungan dan kesehatan yang
terkait dengan penggunaannya. Keprihatinan
publik atas keamanan telah memberikan tekanan
yang luar biasa untuk menilai kembali dampak
toksikologi dan lingkungan dari bahan kimia
sintetik dan protokol yang lebih ketat sedang
243
 dikembangkan (Dayan et al.,
1999b). Cepat tahan herbisida
ecotypes dari gulma juga merupakan ancaman
serius bagi produksi pertanian, dan sejauh ini
272

244
biotipe tahan gulma yang termasuk ke dalam 172
spesies (98 diots dan 64 monokot) terhadap
herbisida telah diidentifikasi (Holt dan LeBaron
1990; Shaner,
1997; Heap, 2003). Bahkan resistansi silang
adalah
cepat berkembang di gulma, di mana mereka
tahan terhadap bahan kimia yang dicoba untuk
pertama kalinya atau milik kelas yang berbeda
secara kimia. Kupatt et al. (1993) berpendapat
bahwa resistensi atau resistensi silang di antara
gulma adalah ancaman serius terhadap industri
herbisida dan praktik manajemen gulma. Karena
semua masalah ini, upaya sedang dilakukan
untuk mencari tahu strategi alternatif untuk
manajemen gulma atau untuk beralih ke praktek
pertanian tradisional input rendah. Manajemen
Gulma Terpadu (IWM), di mana semua opsi yang
meningkatkan produksi dipilih, adalah
pendekatan yang kurang diadopsi karena
berbagai kendala ekologi dan ekonomi dan untuk
menentukan ini lebih banyak penelitian pada
ekologi gulma dan biologi diperlukan (Thill et al .,
1991).
Untuk mencari pestisida dan herisida baru,
umumnya empat pendekatan diikuti di seluruh
dunia (Parry, 1989). Ini adalah
1. skrining acak bahan kimia baru
2. kimia imitasi
3. pendekatan biorasional
4. produk tumbuhan alami
Di antaranya, produk tanaman alami menarik
perhatian karena mereka lebih aman karena paruh
waktu lingkungan yang pendek. Untuk mencari
bahan kimia ini, interaksi allelopathic atau
allelochemicals (produk alami yang bertanggung
jawab untuk membawa fenomena alelopati)
mungkin sangat membantu dalam meningkatkan
produksi tanaman dan mengelola gulma. Faktanya,
bahan-bahan kimia / produk tumbuhan sekunder
membentuk dasar dari allelopathy
- istilah yang diciptakan oleh Molisch (1937).
Kemudian,
khususnya selama 3 dekade terakhir, bekerja secara
rapuh berkembang ke arah ini, dan sekarang ini
merupakan cabang penting dari ilmu terapan
termasuk ekologi kimia. Ini melibatkan pelepasan
kimia / metabolit sekunder dari tanaman atau
mikroba yang mempengaruhi pertumbuhan
tanaman lain, sebagian besar melalui negatif dan
jarang melalui efek positif (Rice, 1984). Untuk
mengelola interaksi alel-patologi gulma dapat
dimanfaatkan langsung atau tidak langsung melalui
penggunaan alelokimia sebagai
alternatif untuk herbisida (Macias et al., 1997,
2001; Kohli et al., 1998a; Duke et al., 1996a,
1997, 2000a, 2002; Caldiz dan Fernandez, 1999;
Dayan dkk., 1999b, 2000; Singh et al., 2001;
Vyvyan, 2002).
II. MENCARI POTENSI INTERAKSI
ALLELOPATIK DALAM
AGROECOSYSTEMS
Dalam agroekosistem tanaman, gulma,
pohon, dan mikroba merupakan komponen
biotik, yang tidak hanya berinteraksi di
antara mereka tetapi juga dengan
lingkungan abiotik. Interaksi alelopati antara
berbagai komponen biotik memiliki potensi
besar dalam meningkatkan produksi
tanaman, keragaman genetik, menjaga
stabilitas ekosistem, konservasi nutrisi, dan
terutama dalam pengelolaan gulma dan
hama (Altieri dan Doll, 1978; Putnam dan
Duke, 1978; Kulit 1983b; Purvis, 1990;
Einhellig, 1996; Swanton dan Murphy, 1996;
Weston, 1996; Kohli et al., 1998a; Anaya,
1999; Chou, 1999; Singh et al., 2001).
Dalam mencari produk alelokimia dari
tumbuhan yang lebih tinggi, Solymosi (1994)
mengeksplorasi 430 spesies Hun- garian untuk
kemampuan pengendalian gulma mereka terhadap
rumput lumbung (Echinochloa crus-galli [L.] P.
Beauv.), domba umum (Chenopodium album L.),
rubah hijau (Setaria) viridis [L.] Beauv.),
semanggi putih (Trifolium repens L.), pigmen
gulma merah (Amaranthus retroflexus L.), gula
bit (Beta vulgaris L.), barley mutiara (Hordeum
distichon L.), prosomillet (Panicum) miliaceum
L.), dan mustard putih (Sinapis alba L.). Untuk
mengekstrak senyawa aktif, 13 pelarut organik
dan air yang berbeda digunakan. Sembilan puluh
dua tanaman dipilih sebagai donor dan studi
menyimpulkan bahwa ekstrak aseton dari 10
tanaman, yaitu, knapweed difus (Centau- rea
diffusa Lam.), Mentimun squirting (Ecballium
elaterium [L.] Rich.), Raksasa hogweed
(Heracleum mantegazzianum Somm . & Levier),
mint pennyroyal (Mentha pulegium L.),
Peucedanum cervaria (L.) Lap., Selfheal (Prunella
vulgaris L.), bunga besar selfheal (P. grandiflora
[L.] Scholl.), Selfheal cut-berdaun ( P. laciniata
[L.] Nath.), Mentega umum (Ranunculus arvensis
L.), sapu umum (Sarothamnus scoparius [L.]
Wimm.), Goldenrod Kanada (Solidago canadensis
L.), dan berbulu halus
woundwort (Stachys germanica L.),
ditemukan cocok untuk aplikasi langsung dalam
pertanian skala kecil karena waktu tinggal mereka
di tanah kurang dari 2 bulan.
245
 Gulma alelopati yang ditemukan di lahan
pertanian
sering mengurangi hasil panen dan
memburuk kualitas mereka, menyebabkan
kerugian keuangan yang berat (Putnam dan
Weston, 1986; Kohli et al., 1998a; Qasem dan
Foy, 2001). Lebih dari 200 spesies gulma
telah diidentifikasi untuk menunjukkan efek
allelopathic pada tanaman lain, terutama
tanaman. Namun, ini memiliki sedikit arti
dalam manajemen gulma, karena gulma
sendiri adalah tanaman yang tidak diinginkan
dan kehadiran / pengenalan mereka di lahan
pertanian dapat menyebabkan lebih banyak
masalah. Namun demikian, properti ini dapat
dimanfaatkan untuk pengelolaan beberapa
gulma yang sangat agresif, khususnya di
lingkungan akuatik dan dengan demikian
mengurangi ketergantungan pada herbisida
sintetik. Sebagai contoh, tambak Kanada
(Elodea canadensis Michaux) dan curly
pondweed (Potamogeton crispus L.) dapat
dihilangkan dari saluran drainase dengan
pengenalan spikerush jarum (Eleocharis
acicularis [L.] Roem. & Schult.) (Yeo dan
Fisher, 1970). Selanjutnya, allelochemicals
dari beberapa spesies liar dapat diekstrak,
dimurnikan, dan digunakan secara langsung
seperti herbisida sintetis. Misalnya, parthenin
dari parthenium ragweed (Parthenium
hysterophorus L.) (Batish et al., 1997b, 2002b)
dan artemisinin dari Artemisia sp. (Duke et al.,
1987; Dayan et al.,1999a; Duke et al., 2000a).
Tidak seperti gulma, tanaman memiliki potensi
besar dalam mengelola gulma dengan berbagai
cara. Mereka dapat secara langsung
mengganggu spesies liar dengan melepaskan
bahan kimia ke lingkungan. Batish et al.
(2001a) telah mendaftarkan 35 tanaman yang
mempengaruhi gulma. Properti ini dapat
dikapitalisasi untuk manajemen gulma
berkelanjutan. Selanjutnya, residu tanaman
seperti rye, bunga matahari, gandum, dan
barley, dll juga bisa sangat berguna dalam
menekan gulma (Kulit, 1983b; Liebl dan
Worsham, 1983; Barnes dan Putnam, 1986;
Weston, 1996; Batish et al .,
2001a). Sifat allelopathic dari cover, smother,
dan tanaman pupuk hijau atau tanaman yang
ditanam dalam rotasi dapat membentuk strategi
lain yang layak dieksploitasi untuk manajemen
gulma (Kulit, 1987; Jordan, 1993; Weston,
1996; Anaya, 1999; Chou, 1999; Singh et al.,
246
2001; Batish et al. , 2002a). Bahkan kultivar
tanaman berbeda dalam kemampuan kompetitif
mereka karena
 
Tingkat alelokimia juga dapat digunakan di bawah
program IWM. Sebagai contoh, beberapa kultivar
gandum, oat, dan barley diketahui mengandung
lebih banyak asam coumaric, asam vanilat, dan
kandungan scopoletin (alelokimia yang dikenal)
dan secara kompetitif lebih baik daripada yang
lain dengan jumlah yang lebih sedikit (Baghestani
et al.,
1999). Kultivar ini dengan persaingan yang lebih
besar
kemampuan demikian dapat dipilih dan digunakan
untuk mengendalikan gulma (Olofsdotter et al.,
2002). Lebih lanjut, genotipe dengan kandungan
alelokimia yang lebih sedikit dapat ditingkatkan
dengan mentransfer sifat-sifat yang mengkodekan
sintesis allelochemicals. Berbagai jenis alelokimia
yang ditemukan di tanaman juga dapat digunakan
secara langsung sebagai herbisida alami, misalnya,
sorgoleone dari Sorghum sp. (Weston dan
Czarnota, 2001). Keragaman alelokimia
diproduksi oleh tanaman tanaman sebagai strategi
pertahanan (Batish et al., 2001a). Sayangnya,
kultivar modern yang menghasilkan tinggi tidak
memiliki potensi ini karena tekanan seleksi yang
berlebihan terhadap peningkatan hasil kualitatif
dan kuantitatif (Lovett et al., 1982). Foley (1999)
menyatakan bahwa kerabat liar kultivar tanaman
harus diidentifikasi, dan gen yang hilang selama
budidaya harus diperkenalkan kembali melalui
teknik genetika molekuler atau metode
konvensional.
Dalam agroekosistem, pohon yang tumbuh di
bawah berbagai
Program wanatani ous dapat memiliki
pengaruh besar pada tanaman karena ukurannya
yang besar dan karakter berumur panjang. Hampir
80 taksa spesies pohon telah diamati untuk
menunjukkan fenomena alelopati (Rizvi et al.,
1999). Sifat alelopatik dari pohon dapat berguna
dalam mengelola gulma, karena mereka dikenal
untuk mengatur perkecambahan dan pertumbuhan
beberapa spesies gulma (Lodhi dan Rice, 1971;
Rizvi et al., 1980; Alsaadawi et al.,
1985; Mohanty dkk., 1994; Babu et al., 1996;
Kohli et al., 1998b). Beberapa alelokimia
spesifik juga telah diidentifikasi dan diisolasi dari
spesies pohon; misalnya, mimosine diketahui
dapat menghambat spesies gulma seperti billy
goat weed (Agera- tum conyzoides L.) dan
 Mimosa (Mimosa pudica L.) (Chou dan Kuo, keanekaragaman yang lebih rendah. Kultivar
1986; Chou, 1993). Demikian juga, cineole, modern yang unggul telah menggantikan varietas
citronellal, dan beberapa monoterpen lain dari unggul. Tidak diragukan lagi ini telah
Eucalyptus sp. memiliki sifat penekan gulma meningkatkan produksi tanaman secara kualitatif
(Kohli et al., 1998b; Singh et al., 2001, dan kuantitatif, tetapi pada biaya keberlanjutan
2002a). Bahkan, cinmethylin, herbisida agroekosistem. Agroekosistem Berkelanjutan
alami, adalah yang mempertahankan basis sumber
telah disiapkan atas dasar kimia cineole. dayanya, mengandalkan sedikit input buatan dari
Ailanthone - senyawa quassinoid luar sistem pertanian, mengelola hama dan
  penyakit melalui mekanisme regulasi dan mampu
diisolasi dari pohon surga (Ailanthus pulih dari gangguan melalui budidaya dan panen
altissima [Miller] Sw.), telah dilaporkan (Gliesmann ,
memiliki sifat herbicidal yang mirip dengan
glifosat dan paraquat (Heisey, 1996). 1998). Sebaliknya, agroekosistem modern
menyiratkan kultivar tanaman yang responsif
Selain tanaman, gulma, dan pepohonan, terhadap kimiawi, residu, dan residu dari bahan
yang sedangkomponen biotik yang terlihat utama kimia sintetik atau yang disebut pelindung tanaman
ajaib. Keanekaragaman hayati adalah minimum
dari agroekosistem, mikroba - organisme yang
dalam sistem ini karena hal lain selain tanaman
tidak terlihat ke mata telanjang - juga memiliki dihilangkan.
efek mendalam pada interaksi alelopatik yang
terjadi di agroekosistem dengan mengubah /
mengubah sifat kualitatif maupun kuantitatif
alelokimia (Blum et. al., 1999; Pellissier dan
Souto, 1999). Mereka juga menguraikan sisa
tanaman dan dengan demikian memfasilitasi
pelepasan allelochemicals (Rice, 1984). Mereka
tidak hanya membantu melepaskan racun, tetapi
beberapa dari mereka juga menghasilkan
allelochemicals, yang memiliki potensi untuk
menekan pertumbuhan spesies tahunan dan
abadi, bahkan pada konsentrasi yang sangat
rendah (Hoagland, 2001). Potensi sejumlah
bahan kimia asal mikroba sebagai herbisida telah
dieksplorasi dan dibahas secara rinci nanti dalam
teks.

III. AKU AKU AKU. POLA

PENGANGKUTAN,
ALLELOPATI, DAN
MANAJEMEN KAIN

Sejak Perang Dunia II, dengan munculnya

bahan kimia sintetis dan mesin yang mahal,
telah terjadi pergeseran pola tanam dari input
rendah tumpang sari / polikultur menjadi
pertanian / monokultur tunggal dengan input
tinggi yang dikomersilkan yang menghasilkan
247
 Pola tanam yang melibatkan rotasi,
tumpang sari dan agroforestry dapat
meningkatkan kinerja tanaman dan menekan
perkecambahan, pembentukan dan potensi
reproduksi, dll. Dari gulma (Vandermeer,
1989; Liebman dan Davis, 2000) Namun,
sendirian itu tidak memadai. Tumpang sari
serta undersing menyebabkan penekanan
gulma. Baumann dkk. (2000) telah
melaporkan bahwa daun bawang (Allium
porrum L.) yang ditumpangsarikan dengan
seledri (A. graveolens L.) menekan gulma
seperti tanah biasa (Senecio vulgaris L.)
sebesar 58% dan meningkatkan hasil panen
sebesar 10% dibandingkan dengan satu-
satunya memotong. Demikian juga, marigold
Meksiko (Tagetes minuta L.) menurunkan
spesies rumput liar yang terkait (Hunter,
1971). Dengan pola tanam yang berbeda /
beragam, gulma menghadapi berbagai faktor
stres dan mortalitas karena spesies tanaman
yang berbeda dan praktik manajemen yang
berbeda yang mengarah pada kondisi yang
tidak ramah bagi mereka. Kedua, beragam
tanaman menghabiskan sumber daya dengan
cepat, meninggalkan sangat sedikit untuk
gulma dan dengan demikian mengurangi
kemungkinan kelangsungan hidup mereka.
Lebih lanjut, sifat allelopathic dari beberapa
tanaman dapat menambahkan beberapa
alelokimia di tanah, yang pada gilirannya
menghilangkan peluang gulma untuk
berkembang. Oleh karena itu, disadari secara
global bahwa untuk menjaga keberlanjutan
lahan pertanian, praktek pertanian tradisional
harus dihidupkan kembali, dan dalam proses
ini alelopati dapat memainkan peran penting,
jika tanaman yang menunjukkan fenomena ini
digunakan secara selektif.
A. Rotasi Tanaman
Rotasi tanaman melibatkan bergantian
tanaman yang berbeda secara sistematis pada
lahan yang sama dan merupakan strategi
penting untuk manajemen gulma. Karena
gulma cenderung tumbuh subur dengan
tanaman dengan persyaratan pertumbuhan
yang sama, praktik budaya untuk tanaman
tertentu juga menguntungkan pendirian dan
pertumbuhan gulma. Pemangkasan satu-
satunya secara terus-menerus pada lahan
yang sama menghasilkan penumpukan
248
spesies gulma tertentu yang memiliki persyaratan
pertumbuhan yang sama dengan tanaman.
Namun, ketika jenis tanaman yang berbeda
dirotasi, perkecambahan dan siklus pertumbuhan
gulma terganggu oleh berbagai variasi dalam
praktik budaya (seperti persiapan lahan, tanggal
penanaman, dan persaingan, dll.) Yang terkait
dengan setiap tanaman.
Rotasi tanaman adalah salah satu praktik
yang sangat tradisional di mana beberapa
tanaman termasuk penutup dan legum atau
tanaman pupuk hijau ditanam dalam rotasi
pendek dengan tanaman utama dan dapat
mengatasi penyakit tanah (de Candolle, 1832).
Ini meningkatkan produksi tanaman dan
mengurangi kejadian gulma karena kombinasi
dari interaksi allelopathic dan kompetitif
(Overland, 1966). Sampai kontrol gulma abad ke-
20 sebagian besar dicapai melalui rotasi
tanaman (Lee, 1995). Liebman dan Dyck (1993)
dan Liebman dan Ohno (1997) telah
menunjukkan bahwa rotasi tanaman adalah
salah satu teknik yang paling kuat untuk
mengurangi bank benih gulma di tanah dan
kerapatan bibit. Meskipun telah sedikit
dipraktekkan selama beberapa dekade terakhir,
sekarang telah muncul kembali untuk
mempertahankan keberlanjutan ladang
(MacHoughton, 1973; Wilson et al., 1986). Kulit
(1982, 1983a; Schreiber, 1992) melaporkan
bahwa rota tion dengan bunga matahari
mengurangi kerapatan gulma berdaun lebar dan
rumput. Manipulasi tanaman rotasi allelopathic
dapat memberikan cara yang efektif dari
manajemen gulma. Sebagai contoh, di bawah
studi yang dikurangi atau tidak ada penelitian,
terjadi penurunan yang cukup besar dalam
populasi rubah raksasa (Seataria faberii Herrm.)
Yang diamati ketika rotasi kedelai-jagung-
gandum allelectic diikuti dari pada jagung saja,
dan yang signifikan penurunan input herbisida
diamati (Schreiber, 1992). Pengamatan serupa
dilakukan sebelumnya oleh Forcella dan
Lindstorm (1988) bahwa kerapatan gulma
berkurang ketika jagung dirotasi dengan kedelai
dengan pengolahan tanah dibandingkan dengan
jagung saja.
Alfalfa (Medicago sativa L.) dapat secara
signifikan
bersaing dengan gulma dan mengurangi insiden
mereka, dan musim panas gulma tidak pernah
menduduki alfalfa yang tumbuh sangat tebal
(Derscheid et al., 1961; Peters dan Linscott, 1988).
Dimasukkannya alfalfa dalam urutan rotasi
tanaman secara signifikan mengurangi kejadian
gulma pada tanaman berikutnya (Entz et al., 1995).
Kemudian, Ominski dkk. (1999) atas dasar survei
yang dilakukan di 117 ladang di Manitoba, Kanada,
menemukan bahwa rotasi dengan alfalfa dapat
secara efektif mengurangi insidensi oat liar (Avena
fatua L.), Kanada thistle (Cirsium arvense [L. ]
Scop., Mustard liar (Brasscia kaber [DC.] LC
Wheeler), dan jerami tidur gulma (Galium aparine
L.) pada tanaman sereal berikutnya.Para pekerja ini
menyimpulkan bahwa memasukkan alfalfa dalam
rotasi tanaman dapat menjadi bagian yang efektif.
manajemen gulma terintegrasi.
Keberhasilan rotasi tanaman untuk menekan
gulma tergantung pada urutan tanaman yang
menciptakan berbagai pola persaingan sumber daya,
gangguan alel-patologis, gangguan tanah, dan
kerusakan mekanis sehingga memberikan
lingkungan yang tidak stabil dan dengan demikian
mencegah pertumbuhan dan pembentukan dari
spesies gulma tertentu (Liebman dan Dyck, 1993;
Liebman dan Davis,
2000). Namun, mempertahankan rotasi tertentu
semata-mata untuk supresi gulma mungkin sulit
dicapai karena pengaruh pasar dan kekuatan
ekonomi (Bond dan Grundy, 2001).
B. Tutup dan Tutupi Tanaman
Tanaman penutup ditanam di antara periode
tanaman biasa dan juga dikenal sebagai tanaman
cabai karena efek membayangi mereka yang
disebabkan oleh tegakan yang tebal dan
pertumbuhan yang cepat (Overland, 1966; Foley,
1999). Mereka tidak hanya ditanam untuk tujuan
panen tetapi juga untuk konservasi tanah dan
kelembapan, peningkatan siklus nutrisi, menurunkan
suhu, memasok HMT dalam keadaan darurat,
perlindungan tanaman komersial dari angin, supresi
gulma, dan peningkatan hasil panen (Barnes and
Putnam, 1983; Worsham,
1989; Weston, 1990; Teasdale, 1993; Swanton dan
Murphy, 1996; Reicosky dan Forcella, 1998;
Gallandt dkk., 1999; Masiunas, 1999). Tanaman
penutup kacang-kacangan dapat menggantikan
nitrogen pupuk dan karena itu menjaga kesuburan
tanah (Hesterman et al., 1992). Worsham (1991)
berpendapat bahwa penggunaan tanaman penutup
bahkan dapat menggantikan input herbisida. Ada
sejumlah mekanisme di mana tanaman penutup
dapat menekan gulma seperti persaingan mereka
untuk cahaya (Teasdale, 1993), kelembaban tersedia
(Mayer dan Hartwig, 1986), merangsang mikroba,
bayangan, perubahan faktor fisik tanah seperti pH,
kapasitas penahan air, suhu, dan aerasi, dll. dan
pelepasan allelochemicals (Elliott et al., 1981; Kulit,
1983b; Liebel dan Worsham 1983; Putnam dan
DeFrank, 1983; Weston et al., 1989; Blum dkk.,
1991; Yenish et al., 1995; Liebman dan Davis,
2000). Selain itu, keberadaan tanaman penutup
menyebabkan kematian benih gulma yang lebih
besar oleh pemangsa yang disukai (Cromar et al.,
1999). Karena sifat-sifat ini, tanaman penutup
249
menemukan penggunaan ekstensif di acids from wheat (Shil- ling et al., 1985; Wu et al.,
 program manajemen gulma terpadu (Altieri dan 2000a), and saponins from alfalfa (Miller, 1983;
Liebman, 1988; Swanton dan Weiss, 1991). Miller et al., 1988)
Sejumlah tanaman digunakan sebagai
tanaman penutup atau tanaman cabul. Ini
termasuk barley (Hordeum vulgare L), jagung
(Zea mays L.), buckwheat (Fagopyrum
esculentum Moench.), Rye (Secale cereale L.),
sorgum (Sorghum spp.), Gandum (Triti-cum
aestivum L.), beludru kacang (Mucuna pruriens
[L.] DC), alfalfa, cengkeh (Trifolium spp. -
semanggi merah T. incarnatum L., subterranean
semanggi T. subterraneum L., semanggi putih T.
repens L., semanggi merah T. pratense L.), vetch
berbulu (Vicia villosa Roth.), ubi jalar (Ipomoea
batatas [L.] Lam.), gerbang mutiara morning
glory (Ipomoea tricolor Cav.), kacang jack
(Canavalia ensiformis [L.] DC), jumbi bean
(Leucaena leucocephala [Lam.] de Wit.), cowpea
(Vigna unguiculata [L.] Walp.), asam
(Tamarindus indica L.), asam liar (Lyciloma
latisiliquum [L.] Benth.), fescue tinggi ( Festuca
arundinacea Schreb.), Merayap merah fescue (F.
rubra L.), ryegrass abadi (Lolium perenne L.),
medis (Medicago spp.) Dan millet foxtail
(Setaria italica [L.] Beauv.) (Gliesmann dan
Garcia, 1979; Ramos dkk., 1983; Smeda dan
Putnam, 1988; Abdul-Rahman dan Habib, 1989;
Einhellig dan Rasmussen, 1989; Anaya dkk.,
1990; Fujii et al., 1994, 1995; Weston, 1996;
Fujii, 1999a; Anaya, 1999; Foley, 1999;
Caamal- Maldonando et al., 2001). Banyak di
antaranya bersifat alpukopatik, misalnya,
buckwheat (Eskelsen and Crabtree, 1995), rye
(Barnes and Putnam, 1983,
1987; Yenish et al., 1995), sorgum (Weston,
1996), gandum (Shilling et al., 1985),
cengkeh (Padi,
1995; Breland, 1996), alfalfa (Miller, 1983,
1996), vetch berbulu dan beludru kacang (Fujii,
1999a), gandum (Shilling et al., 1985; Lodhi et
al., 1987), dan Ipomoea spp. (Anaya, 1999).Most
of the allelopathic cover crops exhibit weed-
suppressing potential through their residues
(Rice, 1995; Weston, 1996; Teasdale, 1998).
These release a number of allelochemicals. For
example, hydroxamic acids from rye, maize,
wheat (Niemeyer, 1988), sorgoleone from
Sorghum sp. (Netzley and Butler, 1986),
tricolorin A from Ipo- moea sp. (Pereda-Miranda
et al., 1993), L-DOPA from velvet bean (Fujii et
al., 1991; Fujii, 1999b), DIMBOA and phenolic
250
besides phenolic acids. Weed seeds either fail phytotoxins, and the production of phytotoxic
to germinate through contact with plant or microbial products.
rhizo- sphere exudate or perish in the Worsham and Blum (1992) performed a se-
presence of allelochemicals (Karssen and ries of experiments in North Carolina with four
Hilhorst, 1992; Creamer et al., 1996).
Cover crops can be used a living plant tissue
or living mulch or residues prepared either
through decomposition or after desiccation
with contact herbicides or freezing (Rice
1995; Weston, 1996; Teasdale, 1998). Living
mulches are generally considered to be more
competitive with weeds because they are
actively growing and can com- pete
efficiently for water, nutrients, and light. A
chemically stunted stand of crownvetch
(Coronilla varia L.) gives a better weed
control than dead rye mulch (Hartwig, 1989).
Teasdale and Daughtry (1993) found that
weed suppression by live hairy vetch residues
was more than that by paraquat desiccated
residues. Therefore, weed control can be
maximized by keeping hairy vetch residues
live for longer period rather than
killing/desiccat- ing. Living plant tissue of
wheat, crimson clover, subterranean clover,
and rye inhibited the emer- gence of weeds
like ivyleaf morning glory (Ipo- moea
hederacea [L.} Jacq.) and redroot pigweed
(Lehman and Blum, 1997). However, if these
were used after desiccation with glyphosate,
only wheat and crimson clover were
inhibitory. The inhibitory effect of debris so
formed was influ- enced by soil moisture
(Lehman and Blum, 1997). Likewise,
subclover cover crops when used as living
mulch under field conditions can efficiently
control weeds such as fall panicum (Panicum
dichotomiflorusm Michx.) and ivyleaf
morning glory without affecting the yield of
corn (Enache and Ilnicki, 1990; Ilnicki and
Enache, 1992).
Putnam et al. (1983) demonstrated that several
cover crops like rye, wheat, sorghum, and
barley suppress weed species such as redroot
pigweed, common purslane (Portulaca
oleracea L.), and common ragweed
(Ambrosia artemisiifolia L.) up to 95% within
30 to 60 days when their residues (either
frozen or desiccated with herbicides) are
placed on the soil. This they attributed to
various physical and chemical attributes like
shading, re- duced temperature, release of
251
 tanaman penutup yaitu. rye, semanggi bawah
tanah, semanggi merah, dan vetch berbulu. Tiga
spesies pigweed (Amaranthus retroflexus, A.
spinosus, dan A. hybridus) dikendalikan oleh 80
hingga 100% hingga
4 minggu setelah tanam menjadi tanaman
penutup mati
rye dan semanggi subterranean, sedangkan vetch
berbulu dan semanggi merah kurang efektif dalam
mengendalikan gulma (Worsham dan Blum,
1992). Di hadapan tanaman penutup seperti
barley, rye, oat, semanggi sub-terranean, dan
triticale (Triticale spp.), Kerapatan dan biomassa
beberapa gulma seperti rumput lumbung, domba
biasa, pigweed redroot, ryegrass, dan sicklepod
(Cassia obtusifolia L. = Senna obtusifolia [L.] Irwin
& Barneby) berkurang (Purvis et al., 1985; Putnam
dan DeFrank, 1983; Brecke dan Shilling 1996).
Caamal-Maldolando dkk. (2001) dievaluasi
penggunaan empat tanaman penutup kacang-
kacangan yaitu. beludru kacang [Mucuna
deeringiana (Bort.) Merr.], kacang Jack, Jumbi
bean, dan asam liar pada pertumbuhan gulma
dalam hal jumlah, biomassa, keragaman, dan
kepentingan relatif mereka sambil menentukan
hasil jagung. Mereka mengamati bahwa semua
tanaman penutup ini mengurangi pertumbuhan
gulma, tetapi biomassa gulma sangat berkurang
hingga mencapai 68% ketika velvetbean
digunakan sebagai tanaman penutup hidup.
Fisk dkk. (2001) menyoroti potensi
tanaman penutup tanaman legum tahunan
yaitu. duri medis (Medicago polymorpha L.)
dan laras medis (M. truncatula Gaertn.) untuk
mengurangi kepadatan dan pertumbuhan
gulma di no-till jagung. Moynihan dkk.
(1996) mengamati bahwa biomassa gulma di
musim gugur berkurang sebesar 65% dengan
pertanaman obat dibandingkan dengan kontrol
tanpa penutup. Pearly gerbang morning glory
umumnya ditanam sebagai tanaman penutup
sebelum disemai atau selama bera di lahan
tebu di Morelos, Meksiko (Anaya et al.,
1990). Ekstrak berair mengurangi
pertumbuhan radik rumput lumbung dan
bayam hijau (Amaranthus leucocarpus S.
Wats.) Dengan lebih dari 55% (Anaya et al.,
1990) dan oleh karena itu dapat digunakan
untuk manajemen gulma praktis di
agroekosistem tradisional. Jordan et al. (1999)
melaporkan bahwa tanaman penutup gandum
mengurangi munculnya beberapa gulma
seperti rumput gudang, ducksalad
(Heteranthera limosa [Sw.] Willd.), Redstem
252
(Ammania coccinea Rottb.), Dan yellow
nutsedge (Cyperus esculentus). L.).
Vetch berbulu adalah tanaman penutup yang
terkenal di Indonesia
AS dan Eropa. Ini memberikan sejumlah
keuntungan dalam agroekosistem. Ini adalah
fiksasi ni- trogen, penambahan cepat
biomassa, pencegahan erosi tanah dan
promosi keseragaman tanah, ameliorasi iklim
mikro, dan di atas semua penindasan gulma
karena efek alelopatnya (Fujii, 2001). Residu
vetch berbulu mengurangi munculnya
common lambsquarters (Teasdale, 1993) dan
gulma solans (White et al., 1989). Johnson et
al. (1993) mengamati bahwa mulsa vetch
berbulu sepenuhnya menghambat gulma di
bawah sistem tanpa olah tanah. Dalam sebuah
studi elaboratif, Fujii (2001) menilai 53
tanaman penutup tanaman (26 leguminous, 19
graminaceous, dan 8 lainnya) untuk aktivitas
allelopathic mereka dan menemukan bahwa
tikus seperti vetch berbulu dan beludru
kacang dan tanaman graminaceous seperti
oat, rye, gandum, dan barley adalah yang
paling menjanjikan. Lebih lanjut, atas dasar
serangkaian percobaan lapangan, ia
menyimpulkan bahwa tanaman penutup
musim gugur seperti bulu vetch, rye, gandum,
oat, dan mustard lebih baik dalam
mengendalikan gulma bila dibandingkan
dengan tanaman penutup musim semi. Dia
lebih lanjut mengamati bahwa vetch berbulu
sangat efektif dalam mengendalikan gulma di
sawah yang ditinggalkan, dan itu lebih baik
daripada milkvetch Cina (Astragalus sinicus
L.) - tanaman penutup yang umum digunakan
di Jepang. Lebih jauh lagi, kontrol gulma
lengkap dapat dicapai dengan aplikasi
langsung dari vetch berbulu ke sawah.
Berdasarkan eksperimen ini, ia
menyimpulkan bahwa vetch berbulu adalah
tanaman penutup yang menjanjikan di sawah
yang ditinggalkan, padang rumput, dan pantai
di bagian tengah dan selatan Jepang, dan efek
penghambatannya terhadap gulma mirip
dengan aplikasi herbisida. .
Rumput cogon (Imperata cylindrica [L.]
Raeuschel) secara efektif ditekan di plot
dengan tanaman penutup dari velvetbean
(Mucuna cochinchinensis [Lour.] A. Chev.)
Dan kudzu tropis (Pueraria phaseoloides
[Roxb.] Benth.) Daripada di plot tanpa
tanaman penutup (Chikoye et al.,
2002). Tanaman penutup ini dapat
sepenuhnya dihapus
rimpang rumput cogon dalam 2 tahun (Chikoye
et al., 2002). Kacang beludru sebagai tanaman
penutup dapat secara efektif mengendalikan
speargrass di fal-lows (Sauerborn, 1999) dan di
ladang jagung (Udensi et al., 1999; Akobundu
et al., 2000) dengan mengurangi fotosintesis,
penuaan cepat, dan mati-punggung (Suaka,
1995), dan untuk pemberantasan lengkap yang
tersisa
gumpalan dapat dihapus dengan penyiangan
tangan. Casini

253
 dan Olivero (2001) menemukan bahwa
polong polong, benih lindi dan ekstrak air,
residu, dan akar dari tiga tanaman penutup
yaitu. kudzu, kacang kedelai, dan kacang
beludru mempengaruhi perkecambahan dan
pertumbuhan bibit rumput cogon (Imperata
brasiliensis Tring.), dan keluar dari kacang ini
adalah tanaman penutup yang paling
menjanjikan untuk mengendalikan rumput
cogon.
Meskipun tanaman penutup telah menjadi pilihan yang
layak untuk pertanian berkelanjutan karena mereka
mengendalikan gulma, namun untuk mencapai manajemen
gulma yang lengkap mereka perlu digabungkan dengan
praktik budaya lainnya seperti penggunaan herbisida.
Meskipun demikian, dosis herbisida yang digunakan sangat
berkurang. (Enache dan Ilnicki, 1990; Mohler dan Teasdale,
1993; Moore et al., 1994; Masiunas et al., 1995; Williams et
al., 1998; Jordan et al.,
1999; Abdin et al., 2000). Selanjutnya, rumput liarTrol
melalui mereka adalah spesies yang spesifik dan bervariasi
dengan jenis tanaman penutup, pengelolaan populasi,
persiapan lahan, dan populasi gulma selain kondisi
lingkungan yang berlaku (Teasdale,
1996; Peachy et al., 1999; Teasdale dan Mohler,2000).
Misalnya, Brandsæter dan Netland (1999) melakukan
serangkaian eksperimen dengan vetch berbulu, semanggi
merah, dan subkaya dan menyimpulkan bahwa vetch
berbulu adalah tanaman musim dingin yang menjanjikan di
musim dingin di bawah kondisi Norwegia.
1. Rye (Secale cereale L.) sebagai Model
Tanaman Penutup
Rye adalah contoh yang sangat baik dari
tanaman penutup yang ditanam secara
ekstensif dalam beberapa sistem tanam
(Schonbeck, 1988; Mangan et al., 1995). Ini
tidak hanya murah tetapi memiliki sejumlah
keunggulan lain seperti bertahan hidup dalam
kondisi yang sangat dingin, pencegahan erosi
tanah, supresi gulma, dan toleransi terhadap
residu atrazin (Kirkland, 1993). Ini menekan
pertumbuhan gulma dengan naungan,
menurunkan suhu tanah, memoderasi
fluktuasi suhu, dan bertindak sebagai
penghalang fisik di samping melepaskan
alelokimia (Shilling et al., 1985; Barnes dan
Putnam, 1987; Lanfranconi et al., 1993;
Teasdale, 1993 ). Namun, itu tergantung pada
pilihan tanaman, urutan urutan rotasi, musim,
situasi geografis, dan praktik manajemen
yang dilakukan. ini
254
 
tanaman penutup ideal untuk sistem produksi
sayuran (Weston, 1990; Masiunas et al., 1995;
Abdul- Baki dkk., 1996; Masiunas, 1999)
Penutup hidup musim semi yang ditanam di
musim semi -membunuh rye menghambat gulma
berdaun lebar seperti domba biasa, crabgrass
besar (Digitaria sanguinalis [L.] Scop.), dan
ragweed umum dan mengurangi total massa
gulma sebesar 93% dibandingkan dengan praktik
tanpa olah tanah. Itu dikaitkan dengan gangguan
alelopati dan atribut fisik dari mulsa rye (Barnes
and Putnam, 1983, 1986). Rye yang ditanam di
musim semi yang ditanam di musim dingin
mengurangi biomassa gulma pada kedelai
(Glycine max Merr.) Sebesar 60 hingga 90%
(Ateh dan Doll, 1996). Shilling dkk. (1995)
melaporkan bahwa rye mulch efektif dapat
menekan sicklepod. Studi oleh Wagner-Riddle
dkk. (1997) telah menunjukkan bahwa
membunuh tanaman rye seminggu lebih awal
dari penanaman hasil kedelai dalam pertumbuhan
kedelai terbaik di bawah kondisi hujan yang
bervariasi dan tanah tekstur berpasir. Residu dari
rye yang ditanam jatuh menekan gulma lebih dari
rye yang ditanam di musim semi (Masiunas,
1999). Smeda dan Weller (1996) telah
menunjukkan bahwa jatuhnya rye spring-killed
(cv. Wheeler) yang ditanam sebagai tanaman
penutup adalah alat manajemen gulma yang
potensial untuk meningkatkan produksi tomat
(Lycopersicon esculentum Mill.). Ini
mengendalikan gulma lebih dari 90% dan
mengurangi kebutuhan akan herbisida. Gulma
berbeda dalam kerentanan mereka terhadap
mulsa tanaman penutup rye. Secara umum,
residu rye lebih beracun untuk dicot tahunan,
cukup beracun untuk rumput tahunan, dan sangat
sedikit untuk yang abadi (Putnam, 1986, 1990;
Masiunas, 1999). Panjang dan tingkat penekanan
gulma oleh rye tergantung pada jumlah biomassa
rye, jenis spesies gulma, kondisi lingkungan, dan
praktik budaya (Smeda dan Weller, 1996;
Masiunas, 1999). Namun, kontrol gulma
serbaguna sering diperlukan dalam sistem
tanaman penutup rye (Masiunas et al., 1995;
Masiunas,
1999). Bellinder dkk. (1996) mengamati
bahwa residu rye menekan munculnya gulma dan
pertumbuhan hingga 12 minggu; Namun, efek
terbesar dari residu rye pada gulma terjadi segera
setelah membunuh (Mohler dan Calloway,
1995). Penipisan gulma berlangsung lebih lama
 jika curah hujan terjadi segera setelah
membunuh rye yang diikuti oleh kondisi yang
lebih kering (Weston,
1996). Kehadiran residu rye dan satu her-
Aplikasi bisida meningkatkan hasil labu
(Cucurbita maxima Duch.) Dan jagung
manis
 
sistem kontrol dengan satu aplikasi
herbisida saja (Galloway dan Weston, 1996).
Prezepiorkowski dan Gorski (1994)
mengamati bahwa residu rye bahkan dapat
menekan perkecambahan dan pertumbuhan
gulma tahan herbisida.
Dari semua atribut rye, efek allelopathic
telah menghasilkan banyak minat dalam
kemungkinan penggunaannya untuk manajemen
gulma. Rye dikenal untuk melepaskan allelochemicals
dari bagian yang hidup serta residu yang menumpuk
di permukaan tanah. Ini telah diidentifikasi sebagai
asam hidrokarbon siklik seperti 2,4-dihidroksi-1,4
(2H) -benzoxazin-3-satu (DIBOA), dan
2 (3H) -benzoxazolinone (BOA) dan kompleks
asam fenolik sederhana seperti β-hydroxybutyric acid
(β-HBA), β-phenyllactic acid (β-PLA), ferulic, p-
hydroxybenzoic, salicylic, o-coumaric, dan asam
suksinat (Patrick, 1971; Chou dan Patrick, 1976;
Barnes et al., 1986; Shilling et al., 1986; Barnes dkk.,
1987; Wojcik-Wojtkowiak dkk., 1990; Pérez dan
Ormeño-Nuñez, 1993; Mwaja dkk., 1995). Dari ini,
BOA berkontribusi lebih terhadap alelopati daripada
DIBOA. Mikroba tanah seperti bakteri Gram-positif
Acinetobacter calcoaceticus dapat biotransformasi
BOA menjadi 2,2′-oxo-1,1′-azobenzone (AZOB)
dalam tanah dan kemudian telah diamati lebih bersifat
penghambatan daripada DIBOA dan BOA (Chase et
al. ., 1991a, b). Dengan demikian, transformasi
mikroba dari senyawa ini di dalam tanah dapat
meningkatkan aktivitas herbisida. Namun, dengan
degradasi senyawa ini toksisitas dari komoditas rye
berkurang (Yenish et al., 1995). Jumlah DIBOA dan
BOA berkorelasi dengan biomas rye dan penindasan
gulma (Smeda dan Weller,
1996; Masiunas, 1999). Toksisitas dari rye dan
isi DIBOA dan BOA juga dipengaruhi oleh rezim
kesuburan dan lingkungan produksi dan berkorelasi
negatif dengan produksi biomassa (Mwaja et al.,
1995). Kandungan allelochemical dan pengendalian
gulma oleh rye bergantung pada bagian tanaman, usia,
dan genotipe rye (Barnes et al., 1987; Mwaja et al.,
1995). Hoffman dkk. (1996a) menemukan bahwa
residu akar rye lebih beracun daripada residu pucuk
dalam menekan pertumbuhan rumput lumbung.
C. Tanaman Pupuk Hijau
Tanaman Pupuk Hijau adalah tanaman yang
dibalik atau dimasukkan ke dalam tanah selagi
hijau atau segera setelah dewasa untuk
memperkaya pertanian.
 
tanah budaya. Saat ini istilah ini juga
digunakan untuk mulsa tanaman / vegetasi yang
dipotong segar atau segera setelah pengeringan.
Pupuk Hijau dan tanaman penutup sering
digunakan secara bergantian seperti pada tanaman
kasus terakhir dimasukkan ke dalam tanah di
kemudian hari. Sejumlah tanaman digunakan
sebagai pupuk hijau, tetapi beberapa legum dan
anggota Brassicaceae banyak digunakan. Ini
dipandang sebagai sarana manajemen gulma yang
efektif (Al-Khatib dan Boydston, 1999).
Tanaman pupuk hijau legum yaitu. kedelai dan
cengkeh (Trifolium spp.) tidak hanya sumber
nitrogen yang efektif tetapi juga mengelola gulma
yang bergantung pada kualitas dan kuantitas residu,
kelembaban tanah, suhu, dan faktor lainnya (Dyck
dan Liebman,
1994; Myers et al., 1994; Ohno et al., 2000;
Thönnissen et al., 2000a, b). Dyck dan Liebman
(1994) mengamati bahwa pelepasan allelochemicals
dari cengkeh (Trifolium spp.) Menekan
pertumbuhan gulma pada tanaman berikutnya.
Penambahan serpihan semanggi merah dan vetch
berbulu menyebabkan penurunan yang signifikan
dalam kemunculan dan berat kering kemuliaan pagi
di pagi hari (Ipomoea lacunosa L.) dibandingkan
dengan mereka yang tidak mengalami puing (White
et al.,
1989). Residu semanggi merah diubah menjadi
tanah mengurangi pertumbuhan bibit (Ohno et
al.,
2000) dan kemampuan kompetitif mustard liar
(Sinapis arvensis L.) (Conklin et al., 2002), dan
menghubungkan ini dengan pelepasan fenolik dari
residu semanggi merah (Ohno dan Doolan,
2001).
Sejumlah spesies Brassicaceae yaitu. mustar putih
atau kuning (Brassica hirta Moench), sawi coklat /
India (B. juncea [L.] Czern.), sawi hitam (B. nigra [L.]
Koch.), petani lapangan (B. campestris L. ), rapeseed /
oilseed rape / canola (B. napus L.), dan cress garden
(Lepidium sativum L.) adalah sumber pupuk hijau
yang sangat baik karena cocok dengan rotasi tanaman
setelah setiap musim dingin gandum atau setelah
tanaman musim panas durasi pendek seperti jagung
manis (Al-Khatib dan Boydston,
1999; Narwal, 1999). Beberapa spesies Brassica
dilaporkan sebagai allelopathic dan menekan gulma
karena kehadiran glucosinolates (Boydston dan Al-
Khatib, 1994; Boydston et al., 1994; Brown dan
Morra, 1996; Vaughan dan Boydston, 1997).
Dedaunan B. napus dimasukkan ke dalam populasi
yang dipadatkan tanah dari markas domba biasa, babi
255
buluh redroot, rumput lumbung, nightshade enzimatik dari jaringan tanaman diubah menjadi
berbulu isothiocyanate, nitril, epithinitriles atau
  oxazolidine-2-thiones, dan ionic thiocyanate,
(Solanum sarrachoides Sendtner), dan longspine terutama allyl-, methyl-, benzylyl, β-
sandbur (Cenchrus longispinus [Hack.] Fern.) phenylethyl- isothiocyantes (Fenwick et al.,
Mirip dengan perlakuan herbisida yang dikenal 1983; Eberlein et al., 1998). Isothiocynates
(Boydston dan Hang, 1995). Jatuh-tanam benih-
adalah
perkecambahan ketika dimasukkan ke dalam tanah
di musim semi-berkurang kepadatan dan biomassa inhibitor kuat dari perkecambahan biji (Fenwick
gulma di kentang (Boydston dan Hang, 1996). et al., 1983; Wolf dkk., 1984; Dale, 1986; Brown
Baik B. napus dan B. hirta mengurangi munculnya dan Morra, 1995) dan dapat digunakan sebagai
gulma seperti gembala (Capsella bursa-pastoris bioherbisida yang menjanjikan atau penekan
[L.] Medic.), Kochia (Kochia scoparia [L.] gulma (Vaughan Dan
Schrad.), Dan rubah hijau pada tanaman dewasa
berikutnya; bagaimanapun, tindakan tepat waktu
dari budaya dan praktek kimia harus digabungkan
(Al-Khatib et al., 1997).
Petersen et al. (2001) mengamati bahwa
turnip-
Mulsa perkosaan yang dimasukkan ke dalam
tanah bebas pertumbuhan mayweed tanpa
aroma (Matricaria inodora L.) dan sowthistle
berduri (Sonchus asper [L.] Hill.) Dan
hubungan ini dengan pelepasan
isothiocyanates dari mulsa. Krishnan dkk.
(1998) melaporkan bahwa mustard rapeseed,
coklat dan putih mengumpulkan ke dalam
tanah pada 20 g berat segar jaringan ke 450 g
tanah mengurangi publik, biomassa, dan
tinggi Kochia, tas gembala, rubah hijau, dan
kembali. mencelupkan pigweed ke dalam
kedelai. Spesies Mustard sebagai pupuk hijau
untuk mengurangi biomassa gulma pada
kedelai sebesar 40% 4 minggu setelah
kemunculannya. Namun, tingkat
penumpukan gulma tergantung pada jenis
Brassica spp., Kultivar, struktur benih, dan
target spesies target (Waddington, 1978;
Boydston dan Al-Khatib,
1994; Al-Khatib dan Boydston, 1999). Jenis
pembohong
lebih bersifat alelopati karena kehadiran lebih
banyak jumlah glukosinolat (Choesin dan
Boener,
1991). Balik, tanaman pupuk hijau Brassica
pengaruh gulma unggulan kecil lebih dari
gulma unggulan besar (Boydston dan Al-
Khatib, 1994).
Efek penghambatan dari Brassicas diskusi
dengan glukosinolat volatil yang dengan
sendirinya bukan phytotoxic, tetapi setelah
256
Boydston, 1997). Jumlah produksi Umumnya,
glukosinolat bervariasi dengan jenis spesies efek residu tanaman untuk manajemen gulma
dan kultivar (Vaughn dan Boydston, 1997; menurun setelah 4 sampai 6 minggu karena
Eberlein et al., 1998). hilangnya massa residu dan kerusakan alelokimia
(Patrick et al., 1963; Kimber, 1973; Smeda dan
Weller, 1996). Pengelolaan gulma melalui sisa
D. Residu Tanaman tanaman sangat efektif di bawah tropi-

Tanaman residu didefinisikan sebagai

tanaman atau bagian-bagiannya yang tersisa
di lapangan untuk dekomposisi setelah telah
dirontokkan atau dipanen (Kumar dan Goh,
2000). Sebelumnya ini dianggap hanya
sebagai limbah, tetapi sekarang karena
kegunaannya mereka dianggap sebagai
sumber daya penting yang dapat membawa
perubahan fisik, kimia, dan biologis yang
signifikan di tanah pertanian setelah
amandemen. Dalam sistem pengolahan lahan
konservasi, retensi residu tanaman
memperbaiki tanah, meningkatkan
penyaringan air, mengurangi tenaga kerja,
dan menekan gulma di bawah beberapa
situasi dengan membawa pergeseran flora
gulma atau secara langsung menekan mereka
(Wilson dan Foy, 1990; Moody, 1995; Jones)
et al., 1999; Kumar and Goh,
2000). Kehadiran sisa tanaman di tanah
permukaan sebagai mulsa menekan gulma
melalui alelopathy dan dengan demikian
mengurangi ketergantungan yang lebih besar
pada herbisida (White et al., 1989; Worsham,
1991; Weston, 1996; Batish et al., 2001a).
Residu Rye adalah contoh yang sangat baik
yang membawa penekanan gulma melalui
alelopati (Shilling et al.,
1986; Masiunas, 1999).
Tanaman residu hadir sebagai mulsa juga
menekan gulma, terutama semusim, melalui
fenomena alelambala. Namun, praktik
manajemen tambahan diperlukan (Burgos
dan Talbert,
1996; Yenish et al., 1996). Beberapa residu
juga
dikenal untuk meningkatkan efisiensi
herbisida (Teasdale et al., 1991); Namun,
banyak tergantung pada sisa tanaman,
penempatannya, kondisi lingkungan, dan
pola tanam. Juga telah diamati bahwa jika
mulsa ditambahkan dari sumber luar, ia dapat
menggunakan efek kontrol gulma yang
memadai (Liebl et al., 1992; Masiunas et al.,
1995; Barker dan Bhowmik, 20
257
cal dan negara beriklim hangat karena (1) mereka tetap
tidak subur, (2) tanaman penutup mereka adalah musim
dingin yang keras, dan (3) penurunan suhu karena mulsa
memiliki masalah yang lebih rendah.
Beberapa penelitian menunjukkan bahwa benih
keciltanaman dan gulma lebih rentan terhadap alelokimia di
bawah kondisi lapangan dan ini dikaitkan dengan rasio
permukaan terhadap volume yang lebih besar,
menghasilkan lebih banyak pemaparan benih semacam itu
untuk allelochemicals. Selanjutnya, allelochemicals
dilepaskan oleh residu tanaman tetap di permukaan atas
tanah di mana biji kecil hadir dibandingkan dengan biji besar
yang ditaburkan secara mendalam (Liebman dan Mohler,
2001) Karena efek allelopathic dari residu tanaman adalah
untuk durasi yang pendek, dan sangat bervariasi kondisi
iklim dan edafik, ini merupakan tantangan besar bagi
manajemen gulma. Namun, perubahan dalam lingkungan
fisik seperti nutrisi, suhu, kelembaban, dan kondisi cahaya
dapat membawa manajemen gulma (Teasdale dan Mohler,
1992).
1. Tepung Gandum
Di antara sisa tanaman, banyak
pekerjaan telah dilakukan mengenai
penggunaan residu gandum untuk
mengendalikan gulma. Di bawah kondisi
laboratorium ekstrak air dari residu
gandum ditemukan menjadi penghambat
terhadap berbagai macam gulma seperti
kemuliaan ivyleaf morning, velvetleaf
(Abutilon theophrasti Medik.), Rumput
lumbung dan pigweed redroot, kemuliaan
pagi diadu, sicklepod, oat liar (Avena
ludoviciana). [L.] Dur.), Gulma lobak
(Rapistrum rugosum [L.] Semua), rumput
paradoks (Phalaris paradoxa L.), dan
sowthistle umum atau tahunan (Sonchus
oleraceus L.) (Steinsiek et al., 1980,
1982; Liebl dan Worsham, 1983;
Rambakudzibga, 1991; Zwain, 1996;
Lehman dan Blum, 1997; Jones et al.,
1999; AlSaadawi, 2001). Studi-studi ini
menunjukkan bahwa allelochemicals
terbawa ke tanah dan mempengaruhi
spesies yang kurus. Namun, efek
allelopathic tergantung pada usia puing,
kandungan air, total C, dan N dan pH
tanah (Lehman dan Blum, 1997). Bahkan
biotipe tahan herbisida dari ryegrass
tahunan (Lolium rigidum Gaud.) Ditekan
oleh ekstrak berair residu gandum (Wu et
258
al., 2001). Tanah yang digabungkan dengan
residu gandum juga ditemukan
 
untuk menghambat redroot rumput pigweed
dan lumbung dan phytotoxicity bertahan
hingga 6 minggu (AlSaadawi, 2001).
Aktivitas allelopathic dari residu gandum
juga telah dilaporkan di bawah rumah hijau
dan kondisi lapangan (Banks dan Robinson,
1980; Sementara dan Murray, 1980;
Muminovic,
1991). Di bawah kondisi lapangan, mulsa
jerami gandum ditemukan untuk menekan
gulma seperti babi gulma redam, bayam
berduri (Amaranthus spinosus L.), dan
kemuliaan pagi yang tinggi (Ipomoea
purpurea [L.] Roth.) Setara dengan
perlakuan herbisida di non- plot muluk
(Thilsted dan Murray, 1980; Banks dan
Robinson, 1980). Worsham (1984)
berpendapat bahwa di bawah sistem tanpa-
olah dengan residu gandum bermulut, gulma
pada tanaman berikutnya dapat ditekan
karena alelopati.
Baru-baru ini, Blum dkk. (2002) mengamati hal itu
residu gandum, baik akar maupun tunas, mengurangi
pertumbuhan gulma berdaun lebar seperti ivyleaf
morning glory, redroot pigweed, dan biang kerang
sida (Sida spinosa L.). Residu akar lebih efektif
daripada residu pucuk (Blum et al., 2002). Penerapan
jerami gandum untuk manajemen gulma juga telah
diperluas hingga hutan tanaman. Ekstrak berair dari
jerami gandum ditekan untuk mengikat gulma merah
raspberry (Rubus idaeus L.) sebesar 44%, dan ini
dikaitkan dengan efek allelopathic di bawah kondisi
lapangan (Jobidon et al., 1989a, b). Sejumlah
allelochemicals seperti DIMBOA (2,4-dihidroksi-7-
metoksi-1,4-benzoksazin-3-satu) dan asam fenolik
seperti syringic, vanili, trans-ferulic, cis-ferulic, trans-
p-coumaric, cis-p-coumaric, dan asam p-
hydroxybenzoic telah diidentifikasi dari gandum
(Guenzi dan McCalla, 1966; Shilling et al., 1985;
Lodhi et al., 1987; Wu et al.,
2000a). Mengingat potensi besar residu gandum
dalam mengendalikan gulma, aksesinya / kultivar dari
seluruh dunia sedang disaring untuk kemampuan
gulma-menekan, dan itu berkorelasi dengan konten
alelokimia (Copaja et al., 1991; Nicol et al., 1992;
Wu et al., 1998, 2000a, 2001).
2. Residu Padi
Bahkan residu beras dalam bentuk tunggul dan
jerami dilaporkan mengurangi insiden gulma (6 hingga 8 minggu) herba. Semua bagian Sorgum
dan dengan demikian dapat dimanfaatkan seperti akar, herba dan benih berkecambah
sebagai strategi yang mungkin untuk melepaskan phytotoinhibitor mengurangi
manajemen gulma. Tanah pertumbuhan rumput dan spesies berdaun lebar
  seperti rubah hijau, beludru, dan pigweed halus
tergabung dengan lalat padi terkontrol (Panasiuk et al.,
alelopatik terkontrol (Cyperus iria L.) mirip 1986; Hoffman et al., 1996b). Panasiuk et al.
dengan herbisida (Lin et al., 1992a, b). Residu
beras (var. Sarjoo 52) tergabung dalam tanah
dengan kedalaman 5 sampai 6 cm
5 ton / ha mengurangi biomassa beras hutan
(Echinochloa colona (L.) Link.), Monarch
redstem (Ammania baccifera L.), A. multiflora
Roxb., Dan bunga daun teluk (Phyllanthus
fraternus Webster) (Khan dan Vaishya, 1992).
Narwal (2000) melaporkan bahwa
penggabungan tanah dari jerami atau jerami
stubble ditambah jerami menurunkan insiden
gulma berdaun lebar dan berumput. Ekstrak
berair yang dibuat dari ini menghambat
pertumbuhan gulma berdaun lebar seperti
bindweed lapangan (Convolvulus arvensis L.)
dan rumput gulma seperti gandum liar dan biji
canarygrass biji kecil (Phalaris minor L.)
(Narwal, 2000). Mattice dkk. (1998)
mengidentifikasi beberapa allelochemcials dari
padi yang ditanam di bawah kondisi banjir dan
dataran tinggi. Dalam kondisi banjir,
allelochemcials suka
3-hydroxybenzoic, 3,4-dihydroxycinnamic, dan
4-hydroxyphenylacetic acid dan dalam kondisi
dataran tinggi 4-hydroxy benzaldehyde, 4-
hydroxy ben-zoic acid, 3-hydroxy-4-methoxy
benzoic acid, 4- hydroxy cinnamic acid telah
diidentifikasi dan semua ini memiliki aktivitas
herbisida (Mattice et al., 1998). Sebelumnya,
sejumlah asam fenolik seperti p-
hydroxybenzoic, vanillic, ferulic, o-hy-droxy
phenylacetic, dan asam syringic telah
diidentifikasi dari residu padi di bawah kondisi
air-log (Chou dan Lin, 1976; Chou et al., 1981).

3. Sisa Sorgum
Putnam dan DeFrank (1983) menemukan bahwa
resitivitas Sorgum mengurangi jumlah dan
biomassa dari krokot umum dan crabgrass halus
(Digitaria ischaemum [Schr.] Muhl.) Di bidang
oleh 70 dan 98%, masing-masing. Tanaman
Sorgum berumur 4 hingga 6 minggu diamati
untuk menekan gulma tanpa merusak legum
besar dan 2-
Herba 4 minggu lebih efektif daripada yang lama
259
(1986) melaporkan bahwa penumpukan Miller,
sorgum mengurangi berat kering rumput 1995), dan dengan demikian dapat digunakan
lumbung, pigweed redroot, dan coklat sebagai herbisida alami (Chung dan Miller, 1995).
kemerah-merahan merah. Sisa sorgum Bagian alfalfa kering udara menghambat
melepaskan sorgoleone, glikosida-dhurrin pertumbuhan gulma dataran rendah (Tsuzuki et al.,
sianogenik, dan sejumlah produk perusak 1999). Aplikasi pelet alfalfa mengurangi
fenolat yang menghasilkan penindasan gulma perkecambahan dan pertumbuhan gulma dataran
(Guenzi dan McCalla, 1966; Nicollier et al., rendah seperti rumput lumbung (Echinochloa
1983; Putnam, oryzicola [Vas.] Vas.), Crabgrass Henry (Digitaria
ciliaris [Retz.] Koeler), variabel flatsedge (Cyperus
1988; Weston et al., 1989; Weston, 1996).
difformis L.) , dan arrowleaved Monochoria
Kembali-
(Monochoria vaginalis [Burm. f.] Kunth.) di sawah
Baru-baru ini, Cheema dan Khaliq (2000)
melakukan serangkaian percobaan di daerah
semi kering di Punjab, Pakistan, untuk
mengeksplorasi penggunaan sifat all-
pathorgic dari sorghum untuk pengendalian
gulma pada gandum beririgasi. Mereka
mengamati bahwa semprotan
'Sorgaab' (ekstrak air tanaman dewasa yang
diperoleh setelah direndam dalam air selama
24 jam) mengurangi kerapatan gulma dan
biomassa sebesar 35 hingga 49% dan
meningkatkan hasil gandum sebesar 10
hingga 21%. Para pekerja ini lebih lanjut
menunjukkan bahwa herba herba dewasa
ketika ditambahkan ke tanah pada 2 sampai 6
Mg / ha mengurangi gulma sebesar 40 hingga
50% dan meningkatkan hasil gandum sebesar
15%. Berdasarkan penelitian, mereka
menyimpulkan bahwa sorgaab dapat
digunakan sebagai inhibitor gulma alami
pada gandum beririgasi.

4. Alfalfa
Residues

Abdul-Rahman dan Habib (1989) mengamati

bahwa akar alfalfa yang terdekomposisi dan
tanahnya yang terkait mengurangi
perkecambahan rumput cogon oleh lebih dari
50% dan panjang semai lebih dari 85%. Selain
itu, campuran alfalfa membusuk dan undecayed
di tanah menghambat produksi tanaman baru
dari rimpang rumput cogon oleh lebih dari 30
dan 42%, masing-masing. Alfalfa yang dicacah
dan ekstrak berair mereka menghambat
perkecambahan dan pertumbuhan bibit
sejumlah gulma seperti domba, pigweed,
beludru, rubah raksasa, rumput curang (Bromus
secalinum L., dan rumput kepiting (Chung dan
260
ladang (Xuan dan Tsuzuki, 2001). Kemudian, Xuan et Moyer dan Huang (1997) mengamati bahwa
al. (2001) menunjukkan bahwa pelet alfalfa diterapkan traktat lentil (Lens culinaris Medic.), oat (Avena
pada tingkat 1 sampai 3 ton / ha menghambat sativa L.), residu canola dan barley beracun untuk
munculnya abunome (Doparium junceum Hamilt.), flixweed (Descurainia sophia [L.] Webb.),
Pim- terowongan palsu (Lindernia pyxidaria L.), stinkweed (Thlaspi arvense L.), dan brom berbulu
waterwort bertangkai panjang (Elatine tianda Schk. halus
var. pedicellata Krylov), spikerush jarum atau ramping
(Eleocharis acicularis Roem et. Schult. var. longiseta
Sven.), dan rotala berbunga-bunga (Rotala indica
Koehne var. uliginosa Koehne) dan mengurangi
kemunculan dan berat kering rumput lumbung.
Mereka berpendapat bahwa pelet alfalfa dapat
digunakan sebagai herbisida alami di sawah. Sejumlah
allelochemicals seperti medicarpin, 4-methoxy
medicarpin, sativan,

Sativan 5-metoksi, canavanine, saponin, dan asam

fenolik seperti ferulic, chlorogenic, dan salicylic acid
telah diidentifikasi dari alfalfa (Miller et al.,

1988; Dornbos et al., 1990; Gorski et al., 1991;

Wyman-Simpson et al., 1991; Waller et al., 1995).

5. Residu Tanaman
Lainnya
Dalam studi rinci, Barker dan Bhowmik
(2001) telah menunjukkan bahwa residu
kacang kedelai (Glycine max Merr.), Bunga
matahari, dan jagung memiliki kapasitas
untuk menekan gulma dan meningkatkan
produktivitas tanaman tomat dan labu musim
panas (Cucumis pepo L. var. Melopepo Alef).
Para pekerja ini mencoba beberapa metode
aplikasi residu seperti menanam tanaman uji
sebagai tanaman penutup di musim
sebelumnya dan membuangnya ke dalam
petak di musim berikutnya atau penempatan
residu impor seperti mulsa atau dimasukkan
ke dalam tanah dan menyimpulkan bahwa
aplikasi impor residu paling efektif dalam
mengendalikan gulma di musim awal yang
didominasi oleh markas domba biasa,
quickweed (Galinsoga parviflora Cav.), dan
mencabut pigweed. Manajemen gulma
tambahan diperlukan hanya ketika tanaman
penutup ditanam dan dibuang di lahan yang
sama dan juga jika ada hujan yang
berlebihan. Studi ini menyarankan pelepasan
allelochemical dari residu sebagai
kemungkinan penyebab supresi gulma.
261
(Bromus tectorum L.) daripada gandum dan
menyimpulkan bahwa residu dari tanaman ini
dapat digunakan untuk manajemen gulma yang
selektif dalam gandum dan mengurangi
ketergantungan pada herbisida. Hall et al. (1982)
melaporkan bahwa residu bunga matahari Rusia
mengurangi kepadatan spesies gulma di musim
panen berikutnya. Jones dkk. (1999) mempelajari
efek residu jelai, gandum, kanola, dan buncis
pada kelangsungan hidup dan produksi bahan
kering sowthistle, rumput paradoxa, oat liar, dan
gulma turnip dan menyimpulkan bahwa semua
empat sisa tanaman menekan gulma dan jelai ini
adalah sebagian besar habitatnya. Barley
diketahui bersifat allelopathic dan mengandung
berbagai alelokimia (Liu dan Lovett, 1993a, b).
Sayangnya, residu tanaman juga diketahui
untuk mempengaruhi hasil tanaman lain atau
tanaman yang sama karena pelepasan
allelochemicals, terutama fenolat selama
dekomposisi (Patrick dan Koch, 1958; Guenzi
dan McCalla, 1962, 1966; Guenzi et al., 1967;
Patrick, 1971). , Cochran et al.,
1977; Vaughn et al., 1983; Waller et al., 1987;
Bradow and Connick, 1990; Thorne et al., 1990;
Nelson, 1996; Wanniarachchi dan Voroney,
1997). Dalam beberapa kasus mereka juga
dikenal efek nodulasi dan fiksasi nitrogen
biologis (Rice, 1984; Putnam dan Weston, 1986;
Heckman dan Kluchinski, 1995). Selain itu, sisa
tanaman, jika dibakar, tidak hanya mengubah
karakteristik tanah tetapi juga efek efisiensi
herbisida (Rule, 1991; Bowerman, 1995). Oleh
karena itu, ketika memilih sisa tanaman untuk
manajemen gulma sejumlah faktor seperti kondisi
iklim, manajemen dan praktik tanam, dan jumlah
residu yang ditempatkan di permukaan tanah
harus dijaga.
IV. PENAPISAN VARIETAS CROP
UNTUK KEKUATAN ALLELOPATIK
Holt dan Lovett (1993) menunjukkan bahwa
sifat all-pathic hadir di aksesi liar sejumlah
tanaman tanaman yang menanamkan mereka
perlawanan terhadap hama dan gulma. Namun,
selama proses budidaya, gen pengkodean untuk
sintesis allelochemicals secara tidak sengaja
terkikis karena pembiakan untuk hasil tinggi dan
pertumbuhan cepat. Namun demikian, gen-gen
ini dapat dihubungkan kembali melalui berbagai
262
perbaikan genetik.
 
program ment, dan tanaman ini dapat bertindak
sebagai alat untuk manajemen gulma biologis
(Ebana et al., 2001). Untuk ini varietas liar dari
mana kultivar modern berasal dapat disaring dan
dipilih. Rice (1984) dalam risalah lengkapnya
Allelopathy telah mengutip sejumlah penelitian yang
dilakukan untuk memilih akses allelopathic dari
sejumlah aksesi yang dikumpulkan dari berbagai
wilayah geografis. Tanaman di mana studi tersebut
telah dilakukan atau sedang dilakukan tercantum
dalam Tabel 1. Karena akses yang dipilih berada
pada tahap yang berbeda dari peningkatan dan asal
vis-à-vis sejumlah karakter lain, ini menunjukkan
bahwa sifat alelopati poligenik dan berkorelasi
lemah dengan hasil dan karakter lainnya
(Olofsdotter et al., 1999). Bahkan di antara tanaman
tanaman saat ini, kultivar berbeda dalam sifat
alelopatiknya dan dengan demikian genotip superior
dapat dipilih untuk pengelolaan gulma terpadu (Wu
et al., 1999; Olofsdotter et al.,
2002). Perbedaan-perbedaan ini tercermin
bahkan pada tingkat varietas (Christensen, 1995),
dan dengan demikian memilih varietas penekan
gulma yang sangat mungkin berguna (Christensen,
1994; Seavers and Wright,
1995; Lemerle et al., 1996; Seavers and Wright,
1997).
Di antara berbagai tanaman yang tercantum dalam
Tabel 1, padi telah dipelajari secara ekstensif untuk
skrining varietas penekan gulma dan peningkatan
beras melalui teknik genetika molekuler. Dari
koleksi 17.279 aksesi beras yang tersedia di USDA-
ARS yang dikumpulkan dari 110 negara, hampir
5000 telah dievaluasi untuk efek penghambatan
mereka terhadap redstem dan lumbung rumput dan
12.000 terhadap ducksalad (Dilday et al., 1989,
1991, 1994, 1998, 2001). Hampir 145 aksesi
telah dilaporkan menunjukkan allelopathy melawan
redstem, 412 melawan ducksalad, dan 94 melawan
rumput lumbung. Atas dasar aksesi allkopatik
kromatogram disebut sebagai 'PI312777' dan
nonelektopik sebagai 'Rexmont' (Dilday et al.,
2001). Suksesi alelopati yang kuat bisa mendukung
tekan campuran gulma (redstem dan disc water
hyssop Bacopa rotundifolia [Michx.] Wettst.) oleh
72 hingga 95% bila dibandingkan dengan kontrol
100% oleh herbisida (Lin et al., 1992a, b). Bekerja
pada beras juga telah dilakukan di bagian lain dunia,
seperti Jepang dan Filipina. Fujii (1992)
mengidentifikasi
24 varietas beras dengan lebih dari 75% reduksi-
   maupun dicot tidak hanya di laboratorium tetapi
tion selada dari 189 disaring untuk tujuan ini. Di juga di bawah kondisi lapangan. Olofsdotter
IRRI (International Rice Research Institute, (2001) dan Ebana et al. (2001) telah
Manila, Filipina) 111 varietas padi diuji terhadap menyarankan bahwa penelitian masa depan pada
rumput lumbung dan 11 ditemukan allelopathic alelopati beras harus fokus pada pemahaman
di musim kering dan basah berdasarkan mekanisme yang terlibat dalam ekspresi
percobaan yang dilakukan di laboratorium serta allelopathic dari kedua agronomi.
di bawah kondisi lapangan (Olofsdotter dan
Navarez, 1996; Olofsdotter et al., 1995). Para
pekerja ini menyimpulkan bahwa upaya bersama
diperlukan untuk mengeksplorasi potensi
varietas padi untuk pengelolaan gulma. Varietas
padi disaring untuk potensi allelopathic dan
manajemen gulma milik daerah yang berbeda
dan berada pada berbagai tahap perbaikan dan
menunjukkan potensi besar untuk tujuan
pemuliaan (Olofsdotter, 1998).
Untuk mengidentifikasi allelochemical dari
beras
plasma nutfah, isolasi yang dipandu bioassay
telah direkomendasikan oleh Rimando et al.
(2001). Menggunakan penanda RFLP, lokus
sifat kuantitatif (QTL) yang terkait dengan
efek allelopathic dari beras yang
diidentifikasi dengan tujuan untuk
meningkatkan stratifikasi plasma nutfah
untuk mengendalikan gulma secara biologis
(Ebana et al., 2001). Para pekerja ini telah
melakukan analisis QTL pada populasi F2
yang timbul dari persilangan antara jenis
garis Indica PI312777 (sangat inhibitory) dan
Japonica cultivar Rexmont (kurang
penghambatan) dan mengidentifikasi tujuh
QTLs yang terletak pada kromosom 1, 3, 5, 6
, 7, 11, dan 12. Salah satu QTL pada
kromosom nomor 6 saja menyumbang 16%
dari variasi fenotipik yang diamati. Pemetaan
QTL juga dilakukan pada 142 galur padi
inbrida dengan menggunakan teknik seeding
ester untuk fenotip (Jensen et al., 2001).
Empat efek utama QTL yang terkait dengan
efek allelopathic terhadap rumput lumbung
telah diidentifikasi pada tiga kromosom (no.
2, 3, dan 8), dan ini secara kolektif
menyumbang sekitar sepertiga dari total
variasi fenotipik dari aktivitas allelopathic.
QTL ini bisa sangat penting dalam perbaikan
genetik sifat alelopati pada padi. Olofsdotter
(2001) telah menyimpulkan bahwa kultivar
padi menunjukkan variasi besar pada
alelopati, dan beberapa varietas memiliki
potensi untuk menekan baik gulma monokotil

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TABLE 1
List of Crop Plants Where Accessions/Varieties Have Been Screened for Allelopathic Trait and Allelochemicals
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256
TABLE 1 (continued)
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257
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258
TABLE 1 (continued)
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259
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260
TABLE 1 (continued)
dan pandangan lingkungan, identifikasi alelokimia yang
bertanggung jawab untuk suplai gulma aktual di bawah
kondisi lapangan selain penilaian efek lingkungan dan
toksikologi mereka, dan kelanjutan kerja pada
pemahaman genetika untuk memiliki lengkap Pilihan yang
dibantu penanda DNA untuk memasukkan gen alelopatik
ke dalam kultivar yang menghasilkan tinggi.
Tidak hanya beras tetapi bahkan aksesi gandumtelah
diperiksa untuk efek alelopati terhadap gulma (Spruell,
1984; Wu et al., 1998; Rizvi et al., 2000; Quader et al.,
2001). Sebanyak 38 kultivar roti gandum dan satu gandum
durum (Triticum durum Desf.) Dievaluasi untuk efek
diferensial mereka terhadap ryegrass tahunan oleh
ekstrak bioassay (Wu et al., 1998). Potensi alelambala
secara langsung berkorelasi dengan konten fenolik. Akses
gandum ‘Tasman’ dan
‘AUS # 18060’ ditemukan sangat allelo-patuh terhadap
ryegrass tahunan dan memiliki tingkat alelokimia yang
lebih tinggi (Wu et al.,2000a, b). Karena gandum bersifat
autotoxic dan memberikan efek merusak pada tanaman
lain, Wu et al. (2001) mempresentasikan paket
manajemen untuk menghindari efek-efek berbahaya dari
gandum dan mengeksploitasi efek alelematiknya hanya
untuk pengendalian gulma. Demikian juga, akses liar dari
Triticum speltoides (Tausch) Gren. ex K. Richt.) telah
diidentifikasi bahwa secara signifikan menghambat oat liar
dan lindung nilai India (Sisymbrium orientale L.) (Hashem
dan Adkins, 1996; Quader et al., 2001).
V. ALLELOCHEMICALS — NATURE’S
OWN HERBICIDES
Ada semakin banyak bukti bahwa allelochemi-
cals atau produk tumbuhan alami yang berasal
dari tanaman yang lebih tinggi / mikroba dapat
menjadi agrokimia yang ideal (Putnam, 1983;
Cutler, 1988; Putnam, 1988; Hoagland, 1990;
Rice 1995; Duke et al., 1996a, b ,
1997; Kutchan, 1997; Macias et al., 1997; Duke
et al., 1998; Kohli et al., 1998a; Cutler, 1999;
Dayan et al., 1999b; Hoagland, 1999; Kohli et al.,
1999; Macias et al., 1999a, b; Duke et al., 2000a;
Macias et al., 2000a, c; Hoagland, 2001; Macias
et al., 2001; Duke et al., 2002; Vyvyan, 2002). Ini
disintesis dalam jumlah besar pada tumbuhan
sebagai
metabolit sekunder dan menunjukkan
keragaman membingungkan dalam alam
kimia (Paiva, 2000; Hadacek, 2002).
Faktanya, alam telah menganugerahkan
tanaman dengan herbisida sendiri untuk
mengkompensasi tahap non-motil mereka.
Berbeda dengan kepercayaan sebelumnya
sebagai senyawa fungsional (Ellis, 1997),
mereka sekarang dikenal memainkan
sejumlah fungsi fisiologis seperti pertahanan
terhadap hama dan penyakit, interaksi teratur
antara faktor biotik dan abiotik, dan
peningkatan reproduksi kebugaran tanaman
(Harborne, 1989; Berenbaum, 1995a, b;
Seigler, 1996; Taiz dan Zeiger, 1998).
Namun, ada sedikit bukti genetik molekuler
untuk menunjukkan asal evolusioner mereka
meskipun bahwa beberapa gen kloning yang
mengkode enzim biosintesis telah menjamur
dan genom dari selada thale (Arabidopsis
thaliana [L.] Heynh.) Telah diperintah
(Fachhini, 1999).
. Meskipun begitu banyak keragaman kimia,
semua-
bahan kimia dapat secara luas dicirikan menjadi
fenolitik dan terpenoid. Mereka dilepaskan
melalui vulkanisasi, eksudasi akar, kematian dan
pembusukan tanaman, dan leachasi dari residu
hidup atau yang membusuk (Rice, 1984; Anaya,
1999). Setelah rilis, allelochemicals terlibat dalam
berbagai proses metabolik (Einhellig, 1985;
Mizutani, 1999; Waller et al., 1999). Beberapa
faktor menentukan toksisitas mereka seperti
konsentrasi, laju fluks, usia dan keadaan
metabolik tanaman, dan iklim yang berlaku dan
kondisi lingkungan (Wyman-Simpson et al.,
1991; Kohli et al., 1993; Wardle et al., 1993 ;
Weidenhamer, 1996; Gallet dan Pellissier, 1997;
Nilsson et al., 1998). Jumlah dan produksi mereka
bervariasi dalam kualitas dan kuantitas dengan
usia, kultivar, organ tanaman, dan waktu tahun
(Argandona et al., 1981; Hanson et al., 1981;
Wyman Simpson et al., 1991; Devi et al., 1997;
Burgos et al., 1999; Cambier et al., 2000).
Einhellig (1996) menunjukkan bahwa baik biotik
(penyakit dan kerusakan serangga dan interaksi
rencana dengan herbivora) dan faktor abiotik
(suhu, jumlah nutrisi, dan kelembaban)
meningkatkan jumlah dan biosintesis
allelochemical pada tumbuhan.
261
 Mengingat potensi alam yang luas
produk tanaman / allelochemicals, Duke et al.
(2000a, b) membahas banyak strategi untuk memilih
mereka sebagai kandidat potensial untuk manajemen
gulma. Mereka menyarankan memilih tanaman
alelambala potensial sebagai sumber bahan kimia ini

262
berdasarkan petunjuk chemoecological,
anatomical, atau ethno-botani. Eksplorasi
alelokimia untuk manajemen gulma telah
disarankan untuk menjadi strategi logis (Devine
et al., 1993). Sebagai contoh, Sorgoleone -
hydroxyquinone - dipilih berdasarkan ekologi
kimia dan aktivitas alelopati (Rimando et al.,
1998), dan artemisinin dan hypericin berdasarkan
petunjuk anatomi (Duke et al. , 1994; Tellez et al.,
1999). Birkett dkk. (2001) juga menekankan
bahwa aspek sinyal allelopathy dapat menjadi
nilai yang sangat besar untuk memanfaatkan
manajemen gulma dengan cara yang praktis.
Produk tanaman alami memiliki sejumlah
keunggulan dibandingkan herbisida sintetis
(Dayan et al., 1999b; Duke et al., 1997, 2000c)
karena mereka:
• biasanya memiliki struktur yang kompleks,
menunjukkan keragaman struktural dan
karenanya merupakan sumber senyawa timbal
yang tak ternilai.
• memiliki lebih banyak pusat kiral - karbon
hibridisasi sp3.
• memiliki berat molekul tinggi tanpa jumlah
halogen rendah (klorin atau bromin) atau atom
berat.
• ramah lingkungan karena menurun dengan
cepat di lingkungan.
• memiliki situs target aksi baru yang berbeda
dari herbisida sintetis - sangat dicari setelah
kualitas oleh agrochemists (Dayan et al.,
1999b; Duke et al., 2002).
Dengan mengingat sifat-sifat ideal alelokimia
ini, beberapa memiliki kemampuan menekan
gulma yang dibahas di bawah ini.
A. Dari Tanaman Yang
Lebih Tinggi
1. Senyawa Terpenoid
Hampir 23.000 senyawa dengan cincin
isoprenoid telah dikarakterisasi dan ratusan
senyawa tersebut ditambahkan setiap tahun
(Sacchettini dan Poulter, 1997). Di antara
terpenoid monoterpena yang mudah menguap
dan seskuiterpena lakton menunjukkan
tingkat aktivitas biologis yang lebih besar.
Monoterpena Volatile — Rizvi dkk. (1988)
mengamati bahwa geraniol pada konsentrasi 3
mM selesai menghambat perkecambahan berduri
bayam tanpa efek pada tomat dan berpendapat
bahwa itu dapat digunakan sebagai herbisida
yang mungkin. Vaughan dan Spencer (1993)
menyaring 18 monoterpen yang mudah menguap
untuk aktivitas mereka melawan gulma tahunan
yang terkait dengan kedelai. Mereka mengamati
bahwa monoterpen dengan atom oksigen dalam
struktur mereka, seperti geraniol dan α-terpineol,
secara selektif menghambat perkecambahan
gulma tahunan seperti ryegrass Italia (Lolium
multiflorum Lam.), Crabgrass besar, pigweed
redroot, dan velvetleaf dan menyimpulkan
bahwa ini dapat digunakan sebagai potensi untuk
mengembangkan herbisida masa depan baru.
Kedua 1,4- dan 1,8-cineoles dilaporkan
menunjukkan toksin
icity against sicklepod and lumbard grass
(Romagni et al., 2000). Belakangan, Singh et al.
(2002a) menemukan bahwa monoterpen cineole
dan citronellol beracun terhadap gulma gulma
(Ageratum conyzoides L.). Perkecambahan,
pertumbuhan bibit, kandungan klorofil, dan
aktivitas pernapasan sangat dipengaruhi oleh
perlakuan baik monoterpen dan cineole lebih
efektif. Bahkan cinfethylin herbisida berdasarkan
kimia cineole telah diproduksi dan ditemukan
menjadi beracun bagi sejumlah gulma dicot dan
monocot (Grayson et al., 1987) (Gambar 1).
Baru-baru ini, Singh et al. (2002c) menguji empat
monoterpena yaitu. sitronelal, sitronelol, cineole,
dan linalool melawan coffeeweed, dan
mengamati bahwa monoterpe tidak hanya
menekan perkecambahan cofeeweed tetapi juga
mempengaruhi metabolisme energi karena
mereka mempengaruhi metabolisme pernafasan.
Di antara monoterpena yang diuji, sitronel diikuti
oleh linalool sangat fitotoksik dan dapat
berfungsi sebagai template untuk sintesis
bioherbisida.
Volumile Essential Oils - Kohli et al. (1998b)
menunjukkan bahwa minyak esensial Eucalyptus
spp. kaya monoterpen volatil memiliki potensi
untuk mengelola parthenium ragweed. Dudai
dkk. (1999) mengamati bahwa minyak atsiri
yang diekstraksi dari usus Alkitab (Origanum
syriacum L.), teh hyssop (Micromeria fruticosa
[L.] Druce), dan serai (Cymbopogon citratus
[Nees] Stapf.) Menghambat perkecambahan
empat spesies gulma. dan bisa digunakan
sebagai bioherbisida. Efeknya, bagaimanapun,
tergantung pada jenis tanah. Baru-baru ini,
263
Singh et al. (2002b) menemukan bahwa
minyak atsiri dari eucalyptus beraroma
lemon (Eucalyptus citriodora Hook.)
Menghambat pertumbuhan gulma seperti
beggarticks berbulu (Bidens pilosa L.), oat
liar, dan

264
 coffeeweed (Cassia occidentalis L.) dan konsentrasi yang sama hanya sedikit atau
memegang janji yang baik untuk manajemen tidak berpengaruh pada gandum. Batish et al.
gulma. Tworkoski (2002) menguji 25 minyak (2002b) lebih lanjut menunjukkan
esensial yang berasal dari tumbuhan untuk fitotoksisitas parthenin terhadap dua spesies
aktivitas herbisida dan menemukan 25 yang kurus - oat liar dan beggarticks berbulu.
berasal dari thyme merah (Thymus vulgaris Tidak hanya pertumbuhan tetapi bahkan
L.), musim panas yang gurih (Satureja jumlah pigmen klorofil juga berkurang. Baru-
hortensis L.), kayu manis (Cinnamomum baru ini, Singh
zeylanicum Blume), dan clove (Syzygium
aromaticum [ L.] Merr. Et Perry) yang paling
beracun, menyebabkan kematian sel karena
kebocoran elektrolit yang cepat pada
dandelesion lepas (Taraxacum officinale
Weber di Wiggers). Selanjutnya, aplikasi 5
hingga 10% dari minyak esensial ini
dikombinasikan dengan adjuvan
menyebabkan kematian markas domba
umum, ragweed umum, dan johnsongrass
dalam 1 hari. Dari empat minyak ini,
aktivitas herbisida minyak kayu manis
adalah maksimum, dan ini dikaitkan dengan
kehadiran eugenol yang merupakan 84%
dari minyak.
Sesquiterpene Lactones — Di antara sesaring-
erpene lakton, parthenin dan artemisinin paling
menjanjikan (Gambar 2).
Artemisinin dari apsintus tahunan (Artemi- annua
L.) adalah inhibitor pertumbuhan yang poten dan
phytotoxin (Duke et al., 1987; Chen and Leather,
1990; Lydon et al., 1997). Ini menghambat
pertumbuhan akar pigweed, krokot umum, dan
velvetleaf oleh
50% pada konsentrasi 33 µM. Tidak hanya
artemisinin tetapi bahkan analog strukturalnya
sangat fitotoksik (Dayan et al., 1999a) dan dapat
dieksploitasi untuk program manajemen gulma.
Parthenin dari parthenium ragweed dan
derivatif yang dimodifikasi secara struktural
menunjukkan phyto-toksisitas, dan beberapa
analog lebih kuat dari phytotoxin parthenin
(Saxena et al.,
1991; Kohli et al., 1993; Batish et al., 1997a,
2001b). Datta dan Saxena (2001) mempelajari
sifat pestisida dan pestisida parthenin dan
beberapa analognya dan menyimpulkan bahwa
beberapa analog yang dimodifikasi secara
struktural dapat menjadi pilihan yang lebih baik
untuk pengembangan pestisida dan herbisida.
Batish et al. (1997b) mengamati bahwa parthenin
adalah fitotoksin selektif yang menghambat
perkecambahan dan pertumbuhan gulma billgoat
(Ageratum conyzoides L.), sedangkan pada

265
 et al. (2002d) melaporkan bahwa parthenin merusak
pertumbuhan gulma billgoat dengan mempengaruhi
kemampuan berfotosografi dan pernapasan,
kandungan protein, dan aktivitas spesifik enzim
seperti pro- enzim dan amilase. Studi ini
menyimpulkan bahwa pengamatan tersebut dapat
menjadi indikator yang berguna dalam membangun
mekanisme aksi parthenin. Tidak hanya gulma darat
tetapi parthenin juga dapat digunakan untuk
mengendalikan gulma air (Pandey, 1996).
Sejumlah lakton sesquiterpene lainnya, seperti
sebagai heliannuoles dari bunga matahari yang
dibudidayakan, bersifat bioaktif dan mungkin
terlibat dalam pertahanan tanaman terhadap gulma
dicot (Macias et al., 1999b, c). Germacranolides -
kategori lain dari seskuiterpen - juga aktif secara
biologis dan beberapa metode untuk meningkatkan
aktivitas mereka telah diusulkan (Macias et al.,
1999a, c). Berdasarkan phototoksisitas sejumlah
sesquiterpenes, termasuk podolactones, dan
guanolide sesquiterpenes, Macias et al. (1999a,
2000a, c) mengusulkan beberapa model herbisida
yang dapat berguna dalam program manajemen
gulma di masa depan. Dehydrozaluzanin C adalah
lactone sesquiterpene lain yang menunjukkan
fitotoksisitas terhadap berbagai spesies monocot dan
dicot dan merupakan kandidat potensial untuk
pengembangan herbisida (Macias et al., 2000b).
Cespedes dkk. (2001) mengamati bahwa tujuh
seskuiterpena lakton (arturin, ovatifolin, dan
turunannya) diisolasi dari Podanthus ovatifolius dan
Podanthus mitiqui mengurangi pertumbuhan semai
dan respirasi gulma monokot dan dicot seperti rigata
Italia, semanggi merah, selada, dan kulit meksiko.
tomato (Physalis ixocarpa Brot. Ex Hornem.).

2. Benzoxazinoids

Tanaman sereal memiliki serangkaian senyawa

benzoxazinoid (asam hidroksamat siklik) sebagai
produk tumbuhan alami dan menunjukkan keragaman
aktivitas biologis, termasuk alelopati (Niemeyer,
1988; Frey et al., 1997; Friebe, 2001). Friebe (2001)
menunjukkan bahwa benzoxazinoids mengendalikan
hama, penyakit, dan gulma dari sereal dan
menurunkan input agrokimia. Beberapa senyawa
benzoxazinoid penting adalah 2,4-dihidroksi-1,4-
benzoksazin-3- satu (DIBOA) dan 2,4-dihidroksi-7-
metoksi-1,4-

266
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FIGURE 1. Some monoterpenes with weed-suppressing ability.

FIGURE 2. Some sesquiterpene lactones with potential use as a herbicide.

267
 benzoxazin-3-one (DIMBOA) yang lazim dalam
gandum, jagung dan rye (Gambar 3). Selain famili 3. Glukosinolat (Isosianat)
Poaceae, benzoxazinones juga ditemukan pada
anggota famili Acanthaceae, Ranunculaceae, dan Ini ditemukan dalam anggota keluarga
Scrophulariaceae (Hartenstein and Sicker, Brassicaceae, Residaceae, dan Capparidaceae
1994). Pada tumbuhan ini terjadi sebagai glikosida sebagai metabolit sekunder (Mizutani, 1999).
dan Mereka disimpan dalam tanaman utuh tetapi
ketika tanaman terluka / rusak enzim sepele
dilepaskan sebagai aglikon oleh aktivitas enzim β-
dikenal sebagai myrosinase (thioglucoside
glukosidase. DIBOA sebagian besar ditemukan pada glucohydrolase; EC 3.2.3.1) di hadapan air
rye, sedangkan DIMBOA pada jagung dan gandum. memotong ikatan glukosa-sulfur glukosinolat
DIBOA setelah hidrolisis enzimatik melepaskan 2- diikuti oleh penataan ulang, yang dihasilkan di
(3H) - benzoxazolinone (BOA), yang menunjukkan pembentukan sianida organik (nitril),
phytotoxicity kuat terhadap velvetleaf (Wolf et al.,
1985). Whitenack dkk. (1988) lebih lanjut
menunjukkan bahwa BOA dapat diubah menjadi
diazoperoxide-2,2′- oxo-1,1′-azobenzene (AZOB),
yang menunjukkan 8 sampai
10 kali lebih banyak aktivitas dibandingkan dengan
BOA. Ini
Senyawa menghambat pertumbuhan bibit dari
sejumlah spesies tanaman, termasuk gulma seperti
rumput lumbung, crabgrass, dan prosomillet (Barnes
and Putnam,
1987; Putnam, 1988). Atas dasar kehadiran senyawa
ini, strategi pengelolaan gulma yang berbeda telah
dikembangkan, misalnya, penggunaan sereal sebagai
mulsa / tutup dan tanaman cabul (Putnam dan
DeFrank, 1983; Shilling et al., 1986). Pérez (1990)
melaporkan bahwa baik DIMBOA dan MBOA
menghambat pertumbuhan akar oat liar sekitar
50% pada konsentrasi 0,7 dan 0,5 mM, masing-
masing
tively, sedangkan itu A. sativa dirangsang. Kemudian,
Pérez dan Ormeño-Nuñez (1991) menemukan bahwa
rye cultivar ‘Forrajero-Baer’ memiliki aktivitas
penekan gulma yang lebih tinggi karena keberadaan
DIBOA pada eksudat akar dibandingkan dengan
kultivar gandum.
‘Andifen’. Selanjutnya, pada kerapatan tanaman,
jarak tanam, dan total produksi biomassa tanaman
yang sama, berat kering oat liar dan gulma berdaun
lebar kurang sebesar 81 dan 94%, masing-masing, di
ladang rye daripada ladang gandum (Pérez dan
Ormeño-Nuñez, 1993) .
268
isothiocyanates, oxazolidinithiones, dan (Heisey, 1990, 1996; Dayan et al., 1999c). Di
thiocynates ionik (SCN-) (Gambar 4). Semua antara mereka, ailanthone adalah phytotoxin
produk ini, terutama isothiocyanate, adalah kuat yang ditemukan di A. altissima (Mergen,
catabodile biocidal (Fenwick et al., 1989; Halkier, 1959) (Gambar 5). Senyawa ini memiliki potensi
1999), dan efek biocidal bersifat sangat kompetitif besar untuk manajemen gulma dan dapat
(O’Callaghan et al., 2000). Brown dan Morra digunakan sebagai spektrum luas pasca-
(1997,
1999) menunjukkan bahwa ada banyak bukti
bahwa senyawa-senyawa ini paling baik untuk
mengelola hama dan gulma yang terbawa oleh
tanah, sehingga secara agronomis signifikan dan
dapat dimasukkan ke dalam praktik pertanian dan
hortikultura saat ini. Sebelumnya penulis ini
menunjukkan bahwa glukosinolat yang
mengandung senyawa dapat digunakan sebagai
bioherbisida karena mereka adalah penghambat biji
yang potensial (Wolf et al., 1984; Dale,
1986; Fenwick dkk., 1989; Brown dan Morra,
1993. 1995, 1999; Vaughn dan Boydston, 1997;
Vaughan, 1999). Ada sinergi yang luar biasa dalam
isothiocyanates meningkatkan aktivitas mereka
(Mizutani, 1999). Hirsutin - isothiocynate dari
yellowcress merayap (Rorippa sylvestris [L.]
Bess.) - Juga hadir di sejumlah tanaman lain,
menunjukkan aktivitas penghambatan kuat
terhadap selada dan bahkan pada 10 ppm
pertumbuhan selada berkurang (Kawabata et al.,
1989 ). Oleszek (1987) melaporkan bahwa volatil
dari delapan cruciferous taxa mengurangi
perkecambahan dan pertumbuhan bibit rumput
lumbung dan menghubungkan pengurangan ini
dengan keberadaan isothiocyanate, nitriles, dan
thiocyanate di dalamnya. Petersen et al. (2001)
mengamati bahwa isothiocyanate (methyl-,
phenylethyl-, benzyl-, allyl-, n-propyl-, dan n-
butyl-) dilepaskan dari mulsa turcek-perkosaan
sangat menekan perkecambahan sowthistle berduri,
mayweed tanpa cela. , pigweed halus (Amaranthus
hybridus L.), rumput lumbung, dan blackgrass
(Alopecurus myosuroides Huds.). Mungkin
isothiocyanate ini berinteraksi dengan biji gulma
dalam larutan tanah dan sebagai uap dalam pori-
pori tanah dan membawa inhibisi (Petersen et al.,
2001).

4. Quassinoids

Ada senyawa alami yang pahit-mencicipi dari

keluarga Simaroubaceae yang telah diisolasi dari
sejumlah tanaman, termasuk Ailanthus altissima
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FIGURE 3. Structure of natural benzoxazinoids.

FIGURE 4. Breakdown of glucosinolates and formation of various products

270
herbisida muncul, yang cepat diinaktivasi dalam
tanah (Lin et al., 1995; Heisey 1996). Ailanthone bila
digunakan pada tingkat 0,5 kg / ha benar-benar
menghambat pertumbuhan pigweed redroot, cress
garden, yellow foxtail (Setaria glauca [L.] P. Beauv.),
Dan rumput lumbung.
Hailanthone - quassinoid lain - ditemukan
untuk memiliki aktivitas spektrum luas ketika
diterapkan sebagai herbisida pasca-atau
preemergent. Pada 0,125 kg / ha itu memberikan
kontrol lengkap dari rubah hijau dan sakit kepala. Ini
menyebabkan pemendekan dan pembengkakan akar
di tef (Eragrostis tef [Zucc.] Trotter), dan efeknya
mirip dengan herbisida trifluralin (Ashton and Craft,
1981; Lin et al.,
1995). Dayan dkk. (1999c) mengamati bahwa
quassinoids seperti chaparrinone, glaucarubolone,
dan holacanthone sangat phytotoxic dibandingkan
dengan quassin, neoquassin, dan picrasin. Ini
menghambat pertumbuhan rumput bengkok yang
merayap (Agrostis stolonifera L.) dan selada pada 10
µM dan membunuh tanaman pada 100 µM. Dari
ketiga senyawa ini, holacanthone paling aktif (Dayan
et al.,
1999c). Para pekerja ini menyimpulkan bahwa substi
tution dari gugus fungsional oxymethylene memiliki
efek yang besar pada aktivitas biologis quassinoids
dan analog struktural lebih baik.
5. Cyanogenic Glycosides
Ini adalah kategori lain dari allelochemicals, yang
sangat mirip dengan glucosinolates. Senyawa-
senyawa ini setelah cedera atau kerusakan
menghasilkan h
 
drogen sianida dan aglycone cyanohydrin oleh
aktivitas β-glucosidase. Sorgum spp. mungkin
adalah contoh terbaik yang melepaskan glikosida
sianogenik dan menekan pertumbuhan tanaman
lain (Weston, 1996). Dhurrin - glikosida sianogen
- Ada di beberapa bagian Sorgum dan
sesudahnya
hidrolisis menghasilkan hidrogen sianida,
glukosa, dan p-hydroxybenzaldehyde (Putnam,
1988; Weston, 1996) (Gambar 6). Namun, ia
memiliki toksisitas mamalia yang tinggi (Putnam,
1988).
6. Saponins
Saponin adalah sekelompok glikosida yang
mengandung bagian polar, yaitu, gula
poliglikosidik yang larut dalam air, melekat pada
lipid steroid atau bagian triterpenoidal. Ini
ditemukan di sejumlah tanaman dan menarik
karena berbagai kegiatan mereka (Oleszek dan
Jurzysta, 1987; Waller et al., 1995; Hoagland et
al., 1996). Alfalfa memproduksi allelopathic
saponins seperti medicagic glycosides, yang
memiliki phytotoxicity selektif (Anaya, 1999).
Campuran saponin, termasuk asam medicagenic
(Gambar 7), hederogenin, asam lecernic, asam
zhanic, dan soyasapogenol B, mengurangi
perkecambahan dan pertumbuhan rumput lumbung
dan rumput curang (Bromus secalinum L.) lebih
dari gandum dan tanaman uji lainnya dan dengan
demikian bisa menjadi sumber manajemen gulma
(Waller et al., 1987, 1993, 1995).
Gorski dkk. (1991) melaporkan bahwa saponin
(garam natrium asam medicagenic) dan
canavanine dari alfalfa menghambat pertumbuhan
radikula
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272
FIGURE 5. Structure of ailanthone (a quassinoid).
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FIGURE 6. Breakdown of Dhurrin, a cyanogenic glycoside present in S. bicolor.

FIGURE 7. Structure of medicagenic acid.

273
 Amaranthus sp. dan spesies Lepidium. Birkett dkk.
(2001) menunjukkan bahwa saponin memiliki
kemampuan mengganggu membran tanaman dan
dengan demikian dapat digunakan sebagai herbisida
potensial.
7. Sorgoleone
Sorgoleone adalah p-benzoquinone memancar
dari akar beberapa spesies Sorgum (Gambar 8).
Ini pertama kali diisolasi oleh Netzley dan Butler
(1986) dari akar eksudat S. bicolor (L.) Moensch.
Einhellig dan Souza (1992) dan Nimbal et al.
(1996a) melaporkan fitotoksisitas sorgoleone pada
sejumlah sistem tanaman, termasuk gulma.
Penghambatan pertumbuhan disebabkan oleh
menghambat fotosintesis (transpor elektron PSII)
dan respirasi (Rasmussen et al., 1992; Einhellig et
al., 1993; Nimbal et al., 1996b; Gonzalez et al.,
1997). Cara kerjanya mirip dengan diuron
jenis herbisida seperti s-triazines, phenyl urea,
dan uracils, dll. (Streibig et al., 1999).
Sorgoleone bila diterapkan pada tingkat 0,6 a.i. kg / ha
pada bibit gulma berumur 14 hari dilaporkan
menyebabkan penurunan yang signifikan dalam
pertumbuhan dan perkembangan gulma, terutama yang
berdaun lebar seperti black nights (Solanum nigrum
L.), pigweed redroot, sicklepod, krokot umum, dan
ragweed. Sepuluh hari setelah penghambatan
pengobatan masih lebih banyak dengan klorosis berat
diikuti oleh nekrosis, dan efeknya mirip dengan
herbisida sintetis seperti triazines, diuron - inhibitor
fotosintesis yang dikenal (Weston dan Czarnota,
2001). Ketika tanah diresapi dengan 10 hingga 80
ppmw dari sorgoleone, pertumbuhan tunas krokot
umum, selada, dan pigweed redroot berkurang
(Weston dan Czarnota, 2001). Para penulis
menyimpulkan bahwa sorgoleone memiliki keduanya
sifat pre dan postemergent ketika diterapkan ke tanah
atau pada dedaunan bibit.
8. Juglone
Juglone (5-hydroxy-1,4-naphthoquinone) yang
ditemukan pada anggota famili kenari
(Juglandaceae) adalah salah satu allelochemical
paling fitotoksik (Willis, 2000) (Gambar 9). Hal ini
diketahui menghambat pertumbuhan beberapa
tanaman pada konsentrasi serendah 1 μM (Rietveld,
1983). Williams dan Hoagland (1982) melaporkan
bahwa juglone pada konsentrasi 1 mM mengurangi
274
perkecambahan velvetleaf, redroot pigweed, dan
biang kerai sida. Pada 1 sampai 3 mM, itu
mengurangi germinal dari akar babi dan beludru
merah dan pertumbuhan akar di rumput
lumbung hingga 90% (Shettel dan Balke, 1983).
Baru-baru ini, Kocacaliskan dan Terzi (2001) telah
melaporkan bahwa ekstrak daun kenari (Juglans
regia L.) dan juglone menghambat pertumbuhan
germinal dan bibit cress kebun dan alfalfa.
Juglone telah dilaporkan sebagai inhibitor
mitokondria yang kuat (Koeppe,
1972). Hejl et al. (1993) menunjukkan bahwa
juglone
mengganggu pertumbuhan tumbuhan tinggi
dengan mengganggu fungsi kloroplas dan
mitokondria.
9. Caffeine
Juglone (5-hydroxy-1,4-naphthoquinone) yang
ditemukan pada anggota famili kenari
(Juglandaceae) adalah salah satu allelochemical
paling fitotoksik (Willis, 2000) (Gambar 9). Hal ini
diketahui menghambat pertumbuhan beberapa
tanaman pada konsentrasi serendah 1 μM
(Rietveld, 1983). Williams dan Hoagland (1982)
melaporkan bahwa juglone pada konsentrasi 1
mM mengurangi perkecambahan velvetleaf,
redroot pigweed, dan biang kerai sida. Pada 1
sampai 3 mM, itu mengurangi germinal dari akar
babi dan beludru merah dan pertumbuhan akar
di rumput lumbung hingga 90% (Shettel dan
Balke, 1983). Baru-baru ini, Kocacaliskan dan
Terzi (2001) telah melaporkan bahwa ekstrak
daun kenari (Juglans regia L.) dan juglone
menghambat pertumbuhan germinal dan bibit
cress kebun dan alfalfa. Juglone telah dilaporkan
sebagai inhibitor mitokondria yang kuat (Koeppe,
1972). Hejl et al. (1993) menunjukkan bahwa
juglone
mengganggu pertumbuhan tumbuhan tinggi
dengan mengganggu fungsi kloroplas dan
mitokondria.
 FIGURE 8. Structure of sorgoleone, an allelochemical from Sorghum sp.
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275
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276
FIGURE 10. Structure of caffeine.
FIGURE 9. Structure of juglone.
 100%, respectively, and opined that it could be Agrostemin — a natural product from
used as a natural herbicide (Rizvi et al., 1981). corn
Caffeine is a selective phytotoxin inhibiting the cockle (Agrostemma githago L.) is
spiny amaranth with no effect on black gram reported to
(Vigna mungo [L.] Hepp.) (Rizvi et al., 1987).
However, Waller et al. (1986) suggested that
based on the persistence of caffeine, weed
management through this compound might not
be a wise strategy.

10. Other Compounds

α-T (α-Terthienyl). Plants in the family

Asteraceae particularly the Tagetes sp. produce a
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number of allelochemicals, including α-T

compounds and polyacetylenes. α-T compounds
are strongly photoactivated and phytoinhibitory
worth exploiting for weed management (Lambert et
al., 1991). At 15 kg/ha spray application, it caused
50% growth inhibi- tion of redroot pigweed and
common lambsquarters. Campbell et al. (1982)
reported that α-T and phenylheptatriyne are
phytotoxic to seedlings of com- mon milkweed
(Asclepias syriaca L.), common lambsquarters,
timothy grass (Phleum pratense L.), and red clover.
In case of α-T, 50% killing of the weeds was
observed at the concentrations ranging from 0.15 to
1.93 ppm. It is reported to have a site of action
near PS II and has been patented as a natural
herbicide (Duke, 1986b). However, due to high
mam- malian toxicity, these may probably have
less poten- tial for weed management. Nevertheless,
these can be structurally or chemically modified.
Lignans are another class of compounds
show-
ing strong phytotoxicity. Rimando et al. (1999)
iso- lated four lignans viz. diayangambin,
epiyangambin, diasesartemin, and epiashantin
from the leaves of Texas Sage (Leucophyllum
frutescens [Berl.] I.M. Johnst). These compounds
exhibited strong phyto- toxicity inhibiting seed
germination of bentgrass (Agrostis stolonifera L.
cv. Penncross) and seedling development of
lettuce. Podophyllotoxin — an aryltetralin lignan
present in leaves of mayapple (Podophyllum
peltatum L.) — is another phytotoxin that is more
toxic against monocots such as Italian ryegrass
than the dicot lettuce (Oliva et al., 2002). All these
lignans have been observed to be potent inhibitors
of mitosis affecting microtubular organi- zation
(Duke et al., 2002).
277
 decrease the incidence of weedy forbs in the pas-
tures when applied at the rate of 1.2 g/ha and is used
for controlling weeds in eastern European countries
such as Yugoslavia (Gajic, 1966, 1973; Gajic et al.,
1976; Jelenic, 1987).
Clerodane (Clerod-14-ene-3α,4β,13ζ-triol) iso-
lated from Viguiera tucumanensis (Hook. et Arn.)
Griseb. inhibited the germination and root growth of
common lambsquarters, Johnson grass (Sorghum
halepense [L.] Pers.), and tall morning glory with no
effect on canola (Vaccarini et al., 1999).
Withanolides — the C28 highly oxygenated er- got
type steroids are generally found in the family
solanaceae. Vaccarini and Bonetto (2000) isolated
withanolides and their analogues from Iochroma
australe Griseb. and observed that these inhibited the
growth of common lambsquarters and Johnson grass
and possess selective herbicidal activity.
L-DOPA (L-3,4-dihydroxyphenylalanine) is the
main phytotoxic compound identified from velvet
bean — an important cover crop, and possess bio-
logical activity. It is exuded from the roots and after
reaching soil it selectively affects plants, in- cluding
broadleaved weeds (Fujii, 1994, 1999b).
Umbelliferone (7-hydroxycoumarin) is found
generally in members of family Apiaceae. In a pot
study, it was observed to reduce the growth of
velvetleaf, pigweed, and prosomillet by 25 to 40%
when applied at 11.2 kg/ha (Shettel and Balke, 1983).
In spite of so many compounds from higher
plants tested for herbicidal activity, only a few have
been marketed. For example, cinmethylin (Cinch®)
based on the chemistry of cineole and mesotrione
(Callisto®) — a triketone class of herbicide, based on
the allelochemical leptospermone obtained from
Californian bottlebrush, Callistemon citrinus (Curtis)
Skeels (Grayson et al., 1987; Mitchell et al., 2001).
Such a low degree of marketing potential of natural
phytotoxins/allelochemicals is due to their poor per-
formance under field conditions compared with labo-
ratory studies. Further, many allelochemicals ex- hibit
rapid dissipation under natural conditions and thus fail
to give desired results.

B. From Microbes

Microbes (such as actinomycetes, saprophytic

microbes, plant pathogens, and even microbial algae)
are one of the excellent and lucrative sources

278
 of phytotoxins that can be used as herbicides. In from the culture filtrates of Streptomyces
this context the term ‘bioherbicide’ is generally toyocaensis and found to be phytotoxic against
used for microbes that are inundatively applied barnyard and crabgrass with no activity toward
to weeds or to allelochemicals/phytotoxins
produced by them, and for fungal pathogens, in
particular, the term ‘mycoherbicides’ is applied.
In fact, mi- crobes and their phytotoxins have
become a source of considerable research in the
recent past, and excellent reviews are available
in the literature that provide an insight into this
(Lynch, 1976; Fischer and Bellus, 1983; Misato
and Yamaguchi,
1984; Cutler, 1986; Omura, 1986; Duke,
1986a,b;
Duke and Lydon, 1987; Cutler, 1988; Heisey et
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al., 1988; Hoagland, 1990; Lynch, 1990; Cutler,

1991; Duke et al., 1991; Strobel et al., 1991;
Devine et al., 1993; Rice, 1995; Cutler, 1995;
Barazani and Friedman, 1999; Cutler, 1999;
Hoagland, 2001; Mallik, 2001; Saxena and
Pandey, 2001). Even arbuscular mycorrhizal
fungi forming symbiotic relationship with most
vascu- lar plants also have a considerable
potential as a broad-spectrum biocontrol agent of
non-host weed species and could provide an
ecological based weed management (Jordan et al.,
2000). In a recent study, Kremer and Souissi
(2001) screened 2000 isolates of rhizobacteria and
observed that 32% of these were cyanogenic
releasing HCN, and reducing growth of barnyard
grass, and thus can be used for biological weed
control. Microbes can be used for weed control in
two ways, viz. utilization of inhibi-
tory/allelopathic interactions of microbes towards
weed, or the use of phytotoxins produced by them
(Hoagland, 2001). However, the use of
phytotoxins is preferred because of the logistics of
storage, appli- cation and formulation,
compatibility, and half-life time. Moreover, with
toxin the chances of spreading to non-target
species do not exist (Duke and Lydon,
1987). In addition, phytotoxins from microbes
are
more efficacious and selective when compared
with those from higher plants.

1. Microbial Phytotoxins

Anisomycin was the first microbial phyto-

toxin to be used as a template for first microbe-
based commercial herbicide. It was discovered
279
turnip and other broadleaved horticultural produced by certain strains of bacterium
plants (Yamada et al., 1972). Later, an Rhizobium japonicum in the soybean nodules. It
analogue of anisomycin viz. methoxyphenone is an effective herbicide at the rate of 0.2 kg/ha
or NK049 (4- methoxy-3-3′- (Anon., 1969). Owens (1973) compared the
dimethylbenzophenone) was chemi- cally herbicidal effect of rhizobitoxine with the
synthesized and used as a commercial her- postemergent herbicides anitrole and
bicide for rice fields (Yamada et al., 1974). metflurazone and concluded that it could be used
The methoxyphenone rapidly degrades into for postemergent control of crab- grass in
metabo- lites nontoxic to rice. On the basis of Kentucky bluegrass (Poa pratensis L.)
anisomycin, there are reports of other
microbial metabolites and their synthetic
analogues being screened for potential
herbicidal property, for example,
moniliformin or Irpexil. However,
unfortunately, due to toxicological concerns
and differences in mode of action of template
and parent compound, none has been
commercialized (Fischer and Bellus,
1983). A number of phytotoxins have been
iden-
tified and isolated from microorganism that
can be potential source for weed
management (Table
2). These can be used either directly or as
tem- plates for analogue synthesis for
structure-activ- ity studies. Trans-4-
aminoproline is a phytotoxic metabolites
isolated from the cell filtrates of Ascochyta
caulina. It is reported to be phytotoxic to a
wide range of plants with maximum inhibi-
tory effect against common lambsquarters
and little against monocots (Evidente et al.,
2000). Based on its phytotoxic activity and
lesser zootoxicity, it is poposed as a
promising mycoherbicide for the
biological control of lambsquarters (Netland
et al., 2001).
However, there are some other phytotoxins
with weed-suppressing ability and have been
pro- posed as herbicides but could not be
commercial- ized because of their toxicity
toward other life forms, including mammals.
These are cercosporin from Cercospora
spp., isocercosporin from Scolicotrichum
graminis, fumonisin B 1 from Fusarium
moniliforme, AAL-toxin from Alterna- ria
alternata f. sp. lycopersici, and australifungin
from Sporormiella australis (Shier and
Abbas,
1999; Hoagland,
2001).
Rhizobitoxine is an allelochemical
280
 TABLE 2
Phytotoxins from the Microbes with Weed Suppressing Ability
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281
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274
TABLE 2 (continued)
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TABLE 2 (continued)

275
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276
TABLE 2 (continued)
 Downloaded by [Central University of Punjab] at 04:35 07 January 2015

TABLE 2 (continued)

277
 Downloaded by [Central University of Punjab] at 04:35 07 January 2015

278
TABLE 2 (continued)
 Downloaded by [Central University of Punjab] at 04:35 07 January 2015

TABLE 2 (continued)

279
 Downloaded by [Central University of Punjab] at 04:35 07 January 2015

280
TABLE 2 (continued)
TABLE 2 (continued)

281
 lawns since at this concentration it affects only
crabgrass and not bluegrass.
Several potential phytotoxins have been iso-
lated even from a single species. For example,
Streptomyces hygroscopicus yields two very ac-
tive potential herbicides, geldanamycin and
nigricin. Both these compounds reduce the
growth of garden cress by 50% at concentrations
as low as 1 and 2 ppm, and completely inhibit its
growth at 3 or 4 ppm. Geldanamycin caused
visible ne- crosis on radicles, while it was not so
in case of nigricin (Heisey and Putnam, 1986).
Later, an- other compound hydantocidin was
isolated from S. hygroscopicus (Nakajima et
al., 1991a). Hydantocidin exhibited a broad
spectrum of her- bicidal activity against a
number of monocot and dicot weeds. It was
found to be more active than bialaphos
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(Nakajima et al., 1991a). In a green- house
experiment at 500 mg/L it is known to affect a
number of monocot weeds like barnyard grass,
blackgrass, large crabgrass, giant foxtail, green
foxtail, johnsongrass and wild oat, and di- cot
weeds such as common cocklebur (Xanthium
strumarium L.), jimsonweed (Haplopappus
pluriflorus [Gray] Hall), lambsquarters, tall morning
glory, black nightshade, redroot pigweed, prickly
sida, common ragweed, velvetleaf and wild
mustard, whereas it completely inhibits
horsenettle (Solanum carolinense L.),
purple nutsedge (Cyperus rotundus L.), and
yellow nutsedge (Nakajima et al., 1991a). Issac
et al. (1991b) reported that a compound
coaristeromycin synthesized by Strep- tomyces
spp. inhibit the growth of barnyard grass and
Johnson grass. Herboxidine — a polyketide from
S. chromofuscus inhibited postemergence growth
of rape, buckwheat, and maize by 90 to
282
92% when applied at the rate of 6.9 mg/m 2
(Miller-Wideman et al., 1992).
In spite of so much diversity and reports on
microbial herbicides, only two have been
commer- cially used, bialaphos and glufosinate
(Abbas and Duke, 1995; Dayan et al., 1999b).
Bialaphos was first isolated from Streptomyces
viridochromogenes and S. hygroscopicus (Bayer
et al., 1972; Kondo et al., 1973). It contains a
unique amino acid phosphinothricin (PPT)
linked to two L-alanyl moieties. By hydrolytic
cleavage of 2-alanyl groups, PPT (L-PPT which
is the natural form) is released within plants
(Omura et al., 1984). The biological activity of
bialaphos is attributed to L-PPT. Rupp et al.
(1977) reported phytotoxicity of L-PPT resulting
in a patent. Bialaphos thus is a pro-herbicide
with no in vitro activity of its own and L-PPT is
an active ingredient (Manderscheid and Wild,
1986; Wild and Ziegler, 1989). It has been
commercialized as glufosinate, which is
ammonium salt of DL-PPT. Glufosinate is mar-
keted under various tradenames such as Basta®,
Challenge®, Herbiace®, Ignite®, and Harvest®,
etc. and is used as a nonselective postemergent
herbicide for controlling annual and perennial
weeds from orchards, vineyards, fallows, and
even in herbicide resistant transgenic crops
(Vasil,
1996), with dicots being more sensitive. PPT has
two remarkable properties viz. metabolic degra-
dation in higher plants and easy translocation in
vascular tissue, which makes it an ideal
nonselec- tive herbicide. It is also a potent
inhibitor of en- zyme glutamine synthetase. Both
fertilization and light are reported to enhance its
toxicity (Duke and Lydon, 1987). Its activity is
faster than glyphosate but lower than paraquat
(Duke, 1986a)
 and is biodegradable in 20 to 30 days (Jobidon, fields;
1991). It has also been used for controlling red
raspberry in forest plantations of black spruce
(Picea mariana [Mill.] BSP) (Jobidon, 1991).
All the raspberry plants got killed with its
application at 1 to 2 kg/ha within 3 weeks of
application.

2. Direct Use of Microbes for Weed

Control

Besides phytotoxins, various pathogenic and

nonpathogenic microbes can also be used directly
for weed control. Several excellent reviews are
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avail- able in this regard (Watson, 1991; Cardina,

1995; Kremer and Kennedy, 1996; Boyetchko,
1999; Morris et al., 1999; Rosskopf et al., 1999;
Müller- Schärer et al., 2000; Charudattan, 2001).
Ascochyta caulina is a pathogenic fungus specific
to common lambsquarters and has been suggested
as a potential mycoherbicide for controlling this
weed (Netland et al., 2001). Sugawara (2001)
reviewed the work done in Japan regarding the
use of microbes like Exserohilum monoceras,
Nimbya scirpicola, and Xanthomonas campestris
against weeds like barn- yard grass (E. crus-galli
and E. oryzicola), and kuroguwai (Eleocharis
kuroguwai Ohwi), and an- nual bluegrass (Poa
annua L.). A phytotoxin
4S,5S(8S,9S)-diepoxy-(3R)(10R)-dihydroxyundeca-
1,6-diene isolated from Alternaria brassicola
caused necrotic lesions and serious damage to
kuroguwai – a serious paddy weed in Japan at the
dose of 10 mg (Matsumoto et al., 1992).
Charudattan (1991) has listed over 100
micro- bial pathogens with potential for weed
control, and most of these belong to
Colletotrichum sp., Fusarium sp., and
Alternaria sp. Recently, Charudattan (2001) in
his comprehensive review has included a number
of pathogens used for weed control at several
sites/crops as classic biocontrol agents and as
inundative/bioherbicide control strat- egy. Despite
the long list, only eight have been approved for
use as a bioherbicides (Isaacson and Charudattan,
1999; Charudattan, 2001). These are

1. Collego® from Colletotrichum

gloeosporioides f. sp. aeschynomene to
control northern jointvetch (Aeschynomene
virginica [L.] BSP) in the rice and soybean
283
2. BioChon from Chondrostereum (Charudattan, 2000). In addition, there are
purpureum for controlling broadleaved constraints regarding environ- mental limitations,
weedy trees like black/wild cherry host-specificity, and mass pro- duction, lack of
(Prunus serotina Ehrh.) in the forests; market, and difficulties in registra- tion, etc.
3. DeVine® from Phytophthora (Hoagland, 2001).
palmivora to Boyette et al. (1999, 2002) have reported that
control milkweed/strangler vine kudzu (Pueraria lobata [Willd.] Ohwi) — an
(Morrenia odorata [Hook. et Arn.] inva-
Lindley) in citrus orchards in Florida;
4. BioMal® from Colletotrichum
gloeosporioides f. sp. malvae to control
round-leaved mallow (Malva pusilla
Sm., and Malva sp.). It has now been
defunct and is being considered for
reregistration as Mallet WP®;
5. Dr. BioSedge® from Puccinia
canaliculata for controlling yellow
nutsedge. However, it is not
commercially available;
6. Hakatak from Colletotrichum
gloeosporioides
for controlling silky Hakea/silky
needlbush (Hakea sericea Schrad. &
J.C. Wendl.). It was provisionally
registered only for one year (1990-91);
however, still preparation of its spore is
available for use (Morris et al., 1999);
7. Stump Out® from Cylindrobasium
leave to
control resprouting of cut trees in tree
plan- tations;
8. Camperico® from Xanthomonas
campestris cv. Poae for controlling
annual bluegrass, Kentucky bluegrass,
and Zoysia/Korean velvet grass
(Zoysia tenuifolia Trin.) in golf courses
and turf grasses.

Out of these eight, two, BioMal® and Dr.

BioSedge®, are not available commercially
due to a lack of technical and economic
considerations. Likewise, BioChon is not
registered and another product EHONTROL
based on Chondrostereum purpureum is
under registration (Charudattan,
2001). However, the development and
marketing
of living organisms as bioherbicides are more
com- plex and difficult when compared with
synthetic herbicides and the pure phytotoxins
as is also evi- dent from the fact that only
eight have been ap- proved and six registered
284
 sive exotic weed in the southern USA could be In order to control them various cultural
controlled with the fungal pathogen Myrothecium prac- tices, including allelopathic plants, use
verrucaria, and in this respect these authors have of catch
got a U.S. patent (Boyette et al., 2001). Abbas et
al. (2001) reported that M. verrucaria produces
large amounts of macrocyclic trichothecenes after
cul- turing. Out of these, attranone B and
trichoverrins A and B exhibit phytotoxicity with
very low mam- malian toxicity, and these could
be used a leads for future development of
analogues with more poten- tial for weed control
(Abbas et al., 2002). Microbes can also be used
for controlling parasitic weeds. It is dealt in detail
in the next section.
Lichens, although not microbes (but their
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com-
ponents), are another source of secondary
metabo- lites, particularly usnic acid and
anthraquinones. These are unique chemicals
exhibiting new mo- lecular target sites and
herbicidal activity (Dayan and Romagni,
2001). Usnic acid inhibits p-
hydroxyphenylpyruvatedioxygenase — an enzyme
of carotenoid biosynthesis. In vitro activity of
usnic acid is found to be superior to any other
inhibitor of this molecular target site. Among
lichen-derived an- thraquinones, emodin, and
rhodocladonic acid are highly active against
seedlings of higher plants, par- ticularly grasses.
Because of their simple chemical nature and easy
synthesis, these compounds are ideal candidates for
the production of new herbicides.

VI. POTENTIAL OF ALLELOPATHY FOR

PARASITIC WEED MANAGEMNENT

Parasitic weeds (such as broomrape and

witch- weed, etc.) are difficult to control and thus
pose a major threat to agriculture. These not only
pro- duce enormous seeds but also have long
seed viability. Depending on the degree of
dependence on the host plants, the parasitic
weeds can be holo- or hemi-parasites. Some
parasitic weeds like witchweed and broomrape
require precondi- tioning and host stimulants for
germination.

A. Potential of Allelopathy, Through

Allelochemicals, and Cultural Practices

285
 and trap crops, and intercropping have been tried.
Chittapur et al. (2001) reviewed the potential of
various allelopathic catch and trap crops for man-
agement of parasitic weeds. Recently, Acharya et al.
(2002) have observed that toria (Brassica
campestris var. toria) planted as catch crop at the
density of 140 plants/m2 significantly reduces seed
bank of Egyptian broomrape (Orobanche
aegyptiaca Pers.) by over 20% compared with fallow
fields. Even the allelochemicals have been seen to
stimulate the seed germination of parasitic weeds
much before the host seed germination, thereby
reducing their seed banks in soil. Some of the
specific studies regarding the control of Striga and
Orobanche are discussed below.
Striga species normally germinate in response
to compounds released from the roots of host plant.
These compounds are present in extremely low
amount, and their isolation and identification are
quite difficult. Some of the natural germina- tion
stimulants identified from the host plants are strigol,
alectrol, sorgolactone, and sorgoleone (Wegmann,
1998) (Figure 11). Strigol was first identified from
the cotton plants. Hsiao et al. (1981) demonstrated
that strigol stimulates the germination of witchweed
by 85 to 100% at con- centrations ranging from 10–4
to 10–6 M.
Several natural synthetic stimulants contain- ing
unsaturated lactone ring enhance the germina- tion of
witchweed (Worsham et al., 1962; Johnson et al.,
1976; Musselman, 1980). Even some cou- marin
type compounds also enhance or stimulate
witchweed germination (Rice, 1984). Netzley et al.
(1988) isolated and identified four p-
benzoquinones, including sorgoleone from the
hydrophobic root exudates of IS 8768 grain sor-
ghum, which enhanced the germination of witch-
weed. Out of these, sorgoleone — a known
allelochemical — was most effective. All these
compounds can be used in soil to trigger the suicidal
germination of witchweed and hence can be used for
its management. Sugimoto et al. (1998) and Welzel et
al. (1999) have made extensive studies on regulatory
signals between host and witchweed and the role of
germination stimu- lants. Besides strigol, four
sesquiterpene lactones sharing structural features of
lactone ring were found to have a germination
stimulatory activity against witchweed seeds
(Fischer et al., 1989).

286
FIGURE 11. Structure of Strigol and Sorgolactone, natural germination stimulants of Striga sp.

287
 Later, Fischer et al. (1990) tested 24 natural and
synthetic sesquilactones against seed
germination of witchweed and concluded that the
spatial ar- rangement of skeleton in
conjunction with a
5-membered ring of lactone is important in
stimu-
lating witchweed seed germination.
Intercropping has also been observed to re-
duce the emergence of purple witchweed com-
pared with sole cropping (Carson, 1989). Oswald
et al. (1996) reported that planting of legumes
like red leucaena (Leucaena diversifolia
[Schldl.] Benth.) and sesban (Sesbania sesban
[L.] Merr.) induced the germination of
witchweed and can be useful for its control in the
fallows in Kenya. Later, Oswald et al. (1998)
suggested that inter- cropping with sweet potato
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effectively controls purple witchweed. Recently,
two leguminous in- tercrops viz. silverleaf
(Desmodium uncinatum DC.) and greenleaf (D.
intortum [Mill.] Urb) desmodium have been
found to very effective in controlling purple
witchweed (Striga hermonthica [Del.] Benth.) in
maize crop (Khan et al., 1997; Pickett, 1999;
Khan et al., 2000). Khan et al. (2000)
observed that intercropping with Desmodium sp.
increased the yield of maize crop from 4 to 5
ton/ha to 7 to 8 ton/ha. Later, allelo- pathic
effects of D. uncinatum were seen to be involved
in stimulating the seed germination of witchweed
causing premature colonization of maize and
also inhibiting haustorial development, thereby
controlling the witchweed (Khan et al.,
2002). Further, the root exudates were found to
contain water-soluble chemicals that acted as a
germination stimulant as well as inhibitors of
288
haustorial development (Khan et al., 2002).
Oswald et al. (2002) evaluated the use of peanut,
beans, yellow gram, and soybean as intercrop in
maize to control Striga sp. and concluded that
these intercops reduced the emergence and sur-
vival of Striga by 40 to 120%, and the yellow
gram (Cicer arietinum L.) is the most useful in-
tercrop. However, they opined that for the com-
plete eradication of Striga, intercropping needs
to be combined with handweeding of mature
plants. Sauerborn (1999) observed that using
legume crops like calopo (Calopogonium
mucunoides Desv.) and tropical kudzu (Pueraria
phaseoloides [Roxb.] Benth.) as a ground cover
during the fallow period reduce the seed bank of
purple witch- weed in the soil by around 50%
and can be used effectively for controlling
parasitic plants. The controlling effect of legume
crops was due to the release of stimulating
exudates from roots caus- ing an induced
ineffective germination and thereby reduced seed
bank in soil.
Orobanche — another parasitic weed is also
difficult to control. In contrast to witchweed,
only two germination stimulants, alectrol
and orobanchol, have been isolated recently
from the host red clover (Yokota et al., 1998). In
addition, different host species have been found
to strongly stimulate the broomrape germination
by more than
90%. However, the structure of these compounds
is unknown. Nonetheless, several haustoria in-
ducing compounds have been identified from the
roots of host plants. These are xenognosin A and
B from exudates of tragacanth (Astragalus
gummifer Labill.) and soyasapogenol B from
roots of Chi-
 nese or sericea lespedzea (Lespedeza cunata suitable host for the matu- ration of
[Dum.] G. Don.) (cf. Qasem and Foy, 2001). parasite. Maximum germination of
Of late, Qasem screened allelopathic effects
of residues of 137 weed species on the germina-
tion, growth, and development of branched
broom- rape growing on the host tomato and
reported that some of the residues from the
weeds stimulated germination, whereas the other
prevented its in- festation on the host. The study
concluded that allelochemcials have a great
potential for exploi- tation as an effective
management of parasitic weeds either by
preventing its germination or destroying its soil
bank or effecting their viability or killing
germinating seeds on the host (cf. Qasem and
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Foy, 2001).
Vail et al. (1990) tested 115 synthetic terpe-
noid compounds having structures similar to one
of the four rings of strigol against branched
broom- rape and found that nine of these
compounds were very effective germination
stimulants. These researchers concluded that
monocyclic compounds with chemical structure
similar to strigol show significant activity.
Likewise, de Luque et al. (2000) tested six
sesquiterpene lactone sharing structural features
with strigol for their effect on the germination of
sunflower broomrape and found that parthenolide
and 3,5-dihydroxydehydrocostus lactone
stimulated the seed germination and the effect
was species specific with no effect on branched,
Egyptian, and bean/scalloped broom- rape (O.
crenata Forssk.).
As in Striga, cultural practices such as inter-
cropping (Al-Menoufi, 1992) and crop rotation
(Krishnamurthy and Rao, 1976; Al-Menoufi,
1991) can also reduce the infestation of the
broom- rape. Intercropping with fenugreek,
Egyptian lu- pine (Lupinus termis Forssk.), and
turnip (Bras- sica rapa L.) is found to
significantly reduce infestation of bean and
branched broomrape in faba bean (Vicia faba L.)
and tomato (Al-Menoufi,
1991, 1992; Al-Menoufi and Adam, 1998).
Linke
et al. (1991) observed that growing of woollypod
vetch (Vicia dasycarpa Ten. subsp. villosa) for
3 years significantly reduced seed bank of bean
broomrape in a lentil-based system.
Krishnamurthy and Chandwani (1975) tested 17
crop plants to serve as trap crops to stimulate
seed germination of broomrape without a
289
 broomrape seeds was obtained with chilli (Capsi-
cum annuum L.), soybean, and moth bean (Vigna
aconitifolia [Jacq.] Maréchal) and concluded that
these could be used as excellent trap crops.
Not only broomrape and witchweed, even
dodder (Cuscuta sp.) can be controlled with allel-
opathy. Abdul-Rahman and Habib (1986) and Habib
and Abdul-Rahman (1988) have reported that
extracts from a number of weed species are effective
in controlling dodder.

2. Control of Parasitic Weeds Through

Microbes

The direct use of microbes can also provide an

excellent control of parasitic weeds like broom- rape
(Orobanche sp.) and witchweed (Striga sp.). Duafala
et al. (1975, 1976) observed that natural populations
of Rhizoctonia solani in the fields reduce the seed
germination of branched broom- rape (O. ramosa L.)
and suggested that R. solani could be used for its
biological control. Fusarium oxysporum f. sp.
orthoceras has a great potential for controlling
sunflower broomrape (O. cumana Wall.) and can be
used as an excellent agent for biological control of
this weed (Thomas et al.,
1998). Bedi and Donchev (1991) and Bedi (1994)
found that organic substrates colonized with F.
oxysporum f. sp. orthoceras and incorporated into
soil reduce the seed germination of sunflower
broomrape and even increased the sunflower yield.
The soil application of F. oxysporum causes the seed
gemination of sunflower broomrape causes the
necrosis of germ tube and causes the rate of
infestation of sunflower roots. This fungus pen-
etrates the seeds of sunflower broomrape and
destroys the seed contents, thereby reducing the seed
bank in soil (Thomas et al., 1999). More recently,
Amsellem et al. (2001) have found that two strains
of Fusarium sp., F. arthrosporioides E4a and F.
oxysporum E1d, are specific to Egyp- tian,
nodding/drooping (O. cernua Loefl.) and branched
broomrape and could be used effec- tively as
mycoherbicide for seed or transplant or soil-drench
treatment of tomato and other vegetable crops in
order to protect them from broomrape. Tomato plant
roots dipped in spore suspension of these strains and
planted in broom-

290
 rape-infested soil were protected for 6 weeks,
and nearly 90% control of broomrape could be
achieved by drenching transplant soil with these
strains (Amsellem et al., 2001). Likewise, there
are reports indicating a reduction in germination
of other parasitic weed — witchweed by the fun-
gal pathogens and their isolates, for example,
Fusarium nygamai (Abbasher and Sauerborn,
1992), F. oxysporum (Citola et al., 1995), and
F. solani (Ahmed et al., 2000). Recently, Zonno
and Vurro (1999) have observed that fungal tox-
ins such as trichothecenes inhibit the germination
of purple witchweed.
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VII. MANAGEMENT OF AQUATIC
WEEDS THROUGH ALLELOPATHY
Weeds are harmful not only in terrestrial en-
vironment but also have considerable impact on
the aquatic ecosystem. They choke aquatic eco-
systems like rivers, streams, ponds, and lakes,
etc. posing problems for aquatic flora and fauna.
Some of the serious aquatic weeds are Salvinia
sp., Hydrilla sp., Eichhornia sp., Potamogeton
sp., Pistia sp., Azolla sp., and Polygonum sp.
The aquatic weeds either compete with each
other or exert allelopathic impact, leading to a
shift in aquatic communities (Gopal and Goel,
1993). However, knowledge in this regard is
very frag- mentary unlike the terrestrial weeds.
Aquatic weeds can be controlled by using al-
lelopathic aquatic plants with greater competitive
ability in order to replace the whole community
or by the direct use of allelochemicals (in either
pure or crude form) for their control (Putnam,
1988; Elakovich, 1989; Elakovich and Wooten,
1989). Szezepanski (1977) emphasized that
allelopathy can be a useful tool for the weed
management in the aquatic environment. Oborn
et al. (1954) ob- served that pondweed
(Potamogeton sp.) can be eliminated from
cultures mixed with needle or slen- der spikerush
or with dwarf arrowhead (Sagittaria subulata [L.]
Buch.). Yeo and Fisher (1970) found that
Canadian pondweed and curly pondweed can be
eliminated from a drainage canal with the intro-
duction of needle spikerush. Later, Frank and
Dechoretz (1980) suggested that allelopathy plays
a significant role in the suppression of pondweed
by spikerush. Szezepanska (1971) reported that
decaying aerial parts of common cattail (Typha
latifolia L.), narrow leaved cattail (T. angustifolia
L.), creeping spikerush (Eleocharis palustris
L.), meadow reed grass (Glyceria aquatica [L.]
Wahle.), common club-rush (Schoenoplectus
lacustris [L.] Palla), and sweet flag (Acorus
calamus L.) killed the seedlings of common reed
(Phragmites com- munis Trin.) at the rate of 0.33
kg/l. Out of these meadow reed grass exhibited
the strongest activity, and the activity of leachates
from its decaying leaves was comparable to
trichloroacetate.
A number of species of spike-rush (Eleocharis
spp.) are reported to inhibit the growth of other
aquatic weeds like duckweed (Lemna minor L.)
and hydrilla (Yeo and Thurston, 1984; Sutton
and Portier, 1989, 1991; Wooten and Elakovich,
1991). These could be used for controlling other
undesir- able aquatic weeds (Ashton et al., 1985).
Elakovich and Wooten (1989) reported that
extracts of fra- grant waterlily (Nymphaea
odorata Ait.), fanwort (Cabomba caroliniana
Gray), Eurasian water milfoil (Myriophyllum
spicatum Fernald), and wildcelery or tape grass
(Vallisneria americana Michx.) severely
inhibited the growth of duck- weed.
Ragweed parthenium — a harmful terres-
trial weed could be useful for controlling aquatic
weeds. Pandey et al. (1993) found that 0.5%
(w/v) extracts of dried leaf and powder of
parthenium killed water hyacinth (Eichhornia
crassipes [Mart.] Solms.) in 1 month. The pres-
ence of phenolic allelochemicals and sesquiter-
pene lactone parthenin in the aqueous extracts
was suggested to be the reason for such an
activity. Later, Pandey (1994) reported that
even the residues of parthenium have a
potential to inhibit the growth of giant salvinia
(Salvinia molesta Mitchell) — another serious
weed of water bodies. Further, sesquiterpene
lactone parthenin was found to inhibit the
growth of several aquatic weeds such as water
hyacinth, giant salvinia, water lettuce (Pistia
stratiotes L.), floating aquatic fern (Azolla
nilotica Decne.), giant duckweed (Spirodella
polyrhiza [L.] Schleid.), duckweed (Lemna
pauciocostata Hegelm.), hydrilla (Hydrilla
verticillata [L.f.] Royle), hornwort/coontail
(Ceratophyllum demersum L.), and bushy
pondweed (Najas
291
 graminea Del.) (Pandey, 1996). The inhibitory
effect of parthenin was associated with decline
in water use, root dysfunction, excessive leak-
age of solute from roots indicating a massive
damage to cellular membrane, a decrease in
dehydrogenase activity, and a reduction of chlo-
rophyll content in leaves. Among phenolic
allelochemicals p-hydroxybenzoic acid was
observed to be highly toxic to all the weeds at
50 ppm concentration (Pandey, 1996).
Saxena (1992) tested three allelopathic
plants, tropical/flute reed (Phragmites karka
[Retz.] Trin.), purging nut (Jatropha curcas L.),
and lantana (Lantana camara L.), for the control
of water hyacinth. Both flute reed and purging
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nut inhibited the growth of newly formed leaves
of water hyacinth. However, with the treatment
of lantana residues a complete inhibition was
no- ticed. The toxic allelochemicals of lantana
first affected roots of hyacinth causing them to
shrink and blacken between 2 to 3 days,
whereas after
4 to 5 days the leaves died. On the basis of this
study, it was suggested that lantana could be
highly useful for controlling water hyacinth
(Saxena, 1992). Later, Saxena (2000) observed
that leachates from twigs and flowers of lantana
could also be utilized for controlling water hya-
cinth. Recently, Kathiresan (2000) found that
dried flowers of broadleaf thyme (Coleus
amboinicus L.) at a concentration of 0.1 g/l
caused injury to water hyacinth plants, and at 40
g/l completely killed water hyacinth.
Since the chemical weed control in aquatic
environment is fraught with dangers of con-
taminating water and adverse affects on the
aquatic biota, the use of biological means such
as microbes or microbial herbicides hold a great
potential (Barreto et al., 2000). Some specific
studies in this regard are the use of native
Mexican fungi (Jimenez and Charudattan, 1998)
and Alternaria sp. (Shabana et al., 1995) for the
control of water hyacinth, and Fusarium
culmorum for the control of hydrilla (Smither-
Kopperl et al., 1998). Joye (1990) observed
that pathogenic isolates from Macrophomina
phaseolina greatly reduced biomass of hydrilla
within 3 to 4 weeks of inoculation and thus
have a great potential for the biological control
of this weed.
292
VIII. ROLE OF MOLECULAR GENETIC
TECHNIQUES FOR IMPROVING CROPS
FOR ALLELOPATHIC TRAITS
Since crops (as cover/smother) may serve as
potential tools for weed management, these can
be considerably improved by conventional breed-
ing and lately by molecular genetical techniques.
Unfortunately, this aspect has got little attention
in the field of allelopathy in spite of the overall
progress made with recombinant technology.
Nevertheless, some attempts to understand the
genetic basis of allelopathy and to locate genes/
genetic markers governing the production of
allelochemicals have been made in the recent
past (Niemeyer and Jerez, 1997; Dilday et al.,
1998; Jensen et al., 2001). Probably, the first
attempt in this direction was made by
Panchuk and Prutenskaya (1973) and
Grodzinsky and Panchuk (1974). These workers
studied the differential phytotoxicity of wheat
grass (Agropyron glaucum [Desf.] Roem.) and
wheat. They reported that residue extracts of
wheat grass were more inhibi- tory toward the
germination of radish (Raphanus sativus L.) and
root growth of garden cress than the residue
extracts of wheat, and if cross is made between
them, F1 hybrids having more characters of wheat
grass are also very inhibitory. On the basis of this
studies, it was established that there is a
possibility of transferring allelopathic traits from
one cultivar/wild relative to the other. Under the
ideal situation, this would lead to a dramatic
reduction on the reliance of herbicides used to
suppress weeds in the agroecosystems (Dilday et
al., 1992; Chavez et al., 1999). However, this is
not an easy task, and a lot of information regard-
ing genetics of crops and their wild relatives is
required. Courtois and Olofsdotter (1998)
observed that the first step in this direction is to
find out the genes responsible for encoding the
synthesis of allelochemicals, the number of genes
involved, their inheritance, nonallelic interactions
among them, and cost of yield with the transfer
of genes from one plant to the other.
Unfortunately, genet- ics is the limiting factor in
the allelopathic stud- ies.
As regards the number of genes controlling
allelopathic trait, mono-, bi-, and polygenic
inherit- ance has been hypothesized. Although
most of the
 workers point out that allelopathic traits are
polygenically inherited (Jansen, 1996). Dunn et al.
(1981) reported that the production of
hydroxamate in maize is both mono- and poly-
genetically con- trolled. Marum et al. (1979), on
the other hand, found that in reed canarygrass
(Phalaris arundinacea L.) there are two genes
(double recessive) controlling the synthesis of
gramine. Niemeyer and Jerez (1997) proposed a
polygenic model for the accumu- lation of
DIMBOA in wheat. Since the putative
allelochemicals exhibit diversity in chemical
nature and are synthesized by several pathways, it
is ex- pected that more than one gene may control
their synthesis (Einhellig, 1996; Olofsdotter et al.,
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1995). Further, as the amount of allelochemicals
changes in response to environmental stress,
environment may consequently affect the
functioning of genes encod- ing the synthesis of
allelochemcials (Einhellig, 1996). Several studies
have established that the allelopathic trait is a
quantitative character, and it is very difficult to
breed such characters. Therefore, molecular ge-
netics employing several DNA recombinant tech-
niques could be extremely useful in improving
the crops for allelopathic traits.
Duke et al. (2001) have proposed two ap-
proaches for crop improvement utilizing
allelopa- thy. These are (1) to enhance existing
allelopathic potential of a particular crop, and (2)
to produce transgenics by inserting foreign genes
encoding for a particular allelochemical. In the
first ap- proach, the use of DNA markers such as
PCR (polymerase chain reaction), RAPD
(random amplified polymorphic DNA), RFLP
(restriction fragment length polymorphism), and
AFLP (am- plified fragment length
polymorphism), etc. to locate quantitative trait
loci (QTLs) on the plant genome are extremely
helpful followed by marked assisted selection.
For example, RFLP technique has been used to
identify QTLs for production of glucosinolates in
rapeseed (Toroser et al., 1995). Duke et al.
(2001) have pointed out that Sorghum spp. can
be improved for the synthesis of sorgoleone by
its roots and efforts are being made to explore
the biosynthetic pathway for the pro- duction of
sorgoleone. Even a differentially ex- pressed
gene associated with the synthesis of sorgoleone
has been identified (Yang et al., 2001). Several
crops have been transgenically produced with
bar or pat genes (from microbes), which are
responsible for providing resistance to PPT and
glufosinate — the two natural herbicides (Lydon
and Duke, 1999).
McCouch and Tanksley (1991) opined out
that with the help of QTL maps points of correla-
tion between markers and phenotype are estab-
lished, then the required points are selected.
Olofsdotter (2001) reported that in the case of
rice QTL mapping was done using 142
recombinant inbred rice lines produced as a
result of a cross between Japonica upland
cultivar IAC 165 (strongly allelopathic) and
Irrigated Indica culti- var CO 39 (weakly
allelopathic). Four main QTLs located on three
chromosomes that accounted for
35% of total phenotypic variation have been
iden- tified (Jensen et al., 2001).
Hydroxamic acid DIMBOA, which is associ-
ated with insect, pest and herbicide resistance
(Frey et al., 1997), is probably synthesized
throughout the life span via two pathways, that
is, either from methoxylated lactam (Tipton et
al.,
1973) or from DIBOA (Zuniga et al., 1990).
Niemeyer and Jerez (1997) investigated the
genes responsible for accumulation of DIMBOA
in wheat. Chromosome 4A and 4B were found to
contain genes for transformation of DIBOA into
DIMBOA and chromosome 5B for transfer of
methoxylated lactam to DIMBOA. Additionally,
a gene present on chromosome 4D was found to
inhibit the accumulation of DIMBOA (Niemeyer
and Jerez, 1997). Another example where mo-
lecular genetics is being used as a tool for crop
improvement is Sorghum.
The genes responsible for the biosynthesis
and release of allelochemicals can be incorpo-
rated in the present day high-yielding cultivars
by genetic manipulations because there is a high
degree of variability of allelochemical
production in different crop cultivars. For
example, the amount of sorgoleone in sorghum
(Nimbal et al., 1996a; Czarnota et al., 2001),
DIMBOA in wheat (Niemeyer, 1988; Copaja et
al., 1991; Nicol et al.,
1992; Quader et al., 2001), gramine in barley
(Hanson et al., 1981; Lovett and Hoult, 1992),
hordenine in barley (Lovett et al., 1994), and
scopoletin in oat (Fay and Duke, 1977). For en-
hancing the weed-suppressing ability of crop cul-
tivars plant microbiological genomic and
proteomic technology can also be utilized
293
(Birkett

294
 et al., 2001). In addition, biotechnological meth-
ods can also enhance the production of some
allelochemicals through manipulations of bio-
chemical pathways for effective weed manage-
ment (Dayan et al., 1999b).
CONCLUSIONS
From the above discussion, it is clear that the
phenomena of allelopathy and alleohemicals/
phytotoxins from higher plants as well as
microbes have a great potential for controlling
noxious weeds including parasitic and aquatic
ones. Allelopthic crop plants grown as cover,
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smother or green ma- nure crops can be
successfully exploited for weed management
when incorporated as a part of the cultural
practices under Integrated Weed Manage- ment
Programmes. Crop cultivars with higher com-
petitive ability vis-à-vis allelochemicals’ content
can be screened and improved through various
molecular genetic approaches. The work in this
direction has already begun, and the cultivars
with higher allelopathic potential and
allelochemical content have been made in rice
and their field trials are under way. Further, the
vast repository of allelochemicals from higher
plants and microbes can serve as bioherbicide or
as templates for the synthesis of new herbicides
based on their chemis- try. Although a lot of
screening efforts have been undertaken, yet little
success has been achieved in this direction
because of the difficulties in their synthesis
because of complexity in structure, cost
effectiveness, mammalian toxicity, poor results in
field trials, rapid degradation besides the intellec-
tual property rights linked with them. Thus,
before allelopathic interactions and/or
allelochemicals become an essential part of weed
management programs much needs to be done.
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