Phytohormones are the chemical messengers produced by certain plant-associated bacteria that play crucial role in different plant-microbe interactions ( Costacurta & Vanderleyden, 1995 ; Sindhu et al ., 2017 ). Production of different phytohormones like IAA, gibberellic acid and cytokinins by the PGPR strains have been reported to alter root architecture, leading to more adsorption of nutrients and promotion of plant growth ( Malik & Sin­dhu, 2011 ; Park et al ., 2015 ; Sindhu et al ., 2017 ). These phytohormones affect seed growth, time of flowering, senescence of leaves and fruits, gene expression, cellular division and growth. In targeted cells, phytohormones also regulate cellular processes, pattern formation, ve­getative and reproductive de­velopment and stress responses.

Indole-3-acetic acid is one of the most common and most studied auxins ( Spaepen et al ., 2007 ). Plant res­ponses to IAA vary from plant to plant in terms of sensitivity. The capacity to produce IAA is wide spread among plant associated bacteria ( Patten & Glick, 1996 ; Kloepper et al ., 2007 ; Mishra et al ., 2010 ; Malik & Sindhu, 2011 ) and the numbers of IAA-producing organisms range as high as 80% of total soil bacteria. IAA is also involved in regulating the expression of important compounds in bacteria such as cAMP and amino acids ( Katsy, 1997 ). IAA production may e­nhance plant growth by enhancing root surface area through which more of the metabolites can be exuded or absorbed as nutrients ( Gaudin et al ., 1994 ).

Indole-3-acetic acid has been reported to stimulate plant growth in lower concentrations and in contrast, if the concentration becomes higher, the effect is re­versed and elongation of root and shoot is inhibited. Natural auxins have modes of action similar to many herbicides that interfere with plant growth such as 2, -4 -dich­lo­rophenoxyacetic acid (2, -4 -D) and 2, -4, -5 -trichlorophenoxyacetic acid (2, -4, -5 -T) ( Patten & Glick, 1996 ). Nine strains of Klebsiella pneumoniae were isolated from rhizosphere of wheat var. Lokwan ( Sachdev et al ., 2009 ) and only six K. pneumoniae strains showed in vitro IAA production. Inoculation of strains K11 and K42 caused significant gain in root length of inoculated moth beans (~ 92.71%) over the control. Pot experiment results indicated that all the six IAA- producing Klebsiella strains significantly increased the root length and shoot height of inoculated wheat seedlings over the control. Serwar & Kremer (1995) reported that auxins produced in high concentrations in the rhizosphere by deleterious rhizobacteria may con­tribute towards reduced root growth of weeds. For example, an Enterobacter taylorae isolate with high auxin-producing potential (72 mg L ^-1 IAA-equivalents) was found to inhibit root growth of field bindweed ( Convolvulus arvensis L.) by 90.5% when combined with 1·10 ^-5 M L-tryptophan in comparison with non-treated control. Suzuki et al . (2003) isolated an IAA low-producing spontaneous mutant of P. fluorescens HP72LI and the colonization ability of strain HP72 on the bentgrass root was found higher than that of mutant HP72LI. Colonization of strain HP72 on the bentgrass root caused root growth reduction, whereas strain HP72LI did not show such growth reduction. The results suggested that IAA production by strain HP72 contribute towards the development of short root systems and take advantage of root colonization.

High amount of IAA production by deleterious rhizobacteria Bradyrhizobium japonicum GD3, isolated from soybean rhizosphere, was found to give suppressive effect on growth of morning glory ( Ipomoea spp.) weed ( Kim & Kremer, 2005 ). Similarly, growth suppressive effect on weed great brome ( Bromus diandrus Roth.) was observed by inoculation of Pseudomonas trivialis strain X33d in a mixture of soil/sand/peat ( Mejri et al ., 2010 ). Bromus diandrus plants inoculated with rhizobacterial strain X33d showed low root biomass, short root systems and low surface area, volume and number of tips. On the other hand, growth promoting effect was observed on most of the crops, especially durum wheat ( Triticum durum Desf.) by inoculation of Pseudomonas trivialis strain X33d strain. This growth suppression effect on great brome weed and growth promotion effect on durum wheat was attributed to production of IAA by P. trivialis strain X33d. Meliani et al . (2017) reported that Pseudomonas fluorescens and Pseudomonas pu­tida produced IAA in vitro , at concentrations of 89 µg mL ^-1 and 116 µg mL ^-1 , respectively. High levels of IAA excretion by P. putida gave consistent effects in enhancing the plant growth and vigor index. Recently, bacterial isolates BWA18 and RWA52 with high IAA production ability (53.80 and 19.18 µg mL ^-1 , respectively), were found to cause growth inhibition of Avena fatua weed and stimulated the growth of wheat at 25, 50 and 75 days of observations under pot house conditions ( Dahiya et al ., 2019 ).

ALA is a key intermediate in the biosynthesis of tetrapyrroles, such as porphyrins, vitamin B12, chlo­rophyll (bacteriochlorophyll) and heme. ALA is a natural photodynamic compound, which is effective as a biodegradable herbicide ( Sasikala et al ., 1994 ; Phour & Sindhu, 2019 ) and it has been reported to cause a stimulating effect on the growth and photosynthesis of crops and vegetables ( Sasaki et al ., 1993 ). In plants, the ALA concentration is strictly controlled at less than 50 nmol g ^-1 fresh weight ( Stobart & Ameen-Bukhari, 1984 ). Herbicidal activity has been reported to increase accumulation of several chlorophyll intermediates, such as protochlorophyllide, protoporphyrin IX and Mg-protoporphyrin IX, when plants are treated with exogenous ALA at relatively high concentrations (5-40 mM). ALA has been applied as a favorable biodegradable herbicide and insecticide, which is harmless to crops, humans and animals ( Beck et al ., 2007 ; Bhowmick & Girotti, 2010 ; Johansson et al ., 2010 ; Kang et al ., 2012 ).

Liu et al . (2005) selected, from 36 photosynthetic bacterial strains, seven strains belonging to Rhodopseu­domonas sp.; among them, ˈ99-28ˈ showed the highest ALA production ability. However, herbicidal activity of ALA on several plants has been reported to differ by the application methods. At low concentrations (0.01-10 mg L ^-1 ), ALA showed growth-promoting effects on yield of several crops ( Hotta et al ., 1997 ), whereas it suppressed plant growth at higher concentrations (> 2 mM). Zhang et al . (2006) reported that ALA at low concentrations of 0.3-3 mg L ^-1 promoted development and growth of potato microtubers in vitro , and enhanced protective functions against oxidative stresses, but application of ALA at 30 mg L ^-1 and higher concentrations may induce oxidative damage. Hyun & Song (2007) reported production of IAA and ALA by Rhodopseudomonas strains, which promoted the seed germination and growth of tomato plants under axenic conditions. Chaudhary & Sindhu (2016) found that out of 55 rhizobacterial isolates, only six isolates (HCS7, HCS19, HFS7, HFS9, HFS10 and HFS12) showed ALA production varying from 1.3 to

7.0 µg mL ^-1 . Khandelwal et al . (2018) reported that 80% of the rhizobacterial isolates from the rhizosphere of wheat and mustard produced ALA. More ALA production (> 11 µg mL ^-1 ) was ob­served in eight bacterial isolates. Other 54 isolates produced ALA ranging from 5 to

11 µg mL ^-1 and nineteen isolates lacked ALA production ability. Phour & Sindhu (2019) reported significant reduction (92%) in RDW and SDW of Lathyrus aphaca weed by inoculation of ALA-producing Bacillus flexus strain JIM24 under pot house conditions.

Cyanide production is considered as a major trait of rhizobacteria for biological control of weeds ( Kremer & Souissi, 2001 ), because of its ability to inhibit root cell metabolism and effective inhibition of the cytochrome oxidase pathway. The HCN production has been found to be a common trait of Pseudomonas (88.89%) and Bacillus (50%) in the rhizospheric soil and plant root nodules ( Ahemad & Khan, 2009 ). Owen & Zdor (2001) reported that two strains of cyanogenic rhizobacteria ( Pseudomonas putida and Acidovorax delafieldii ), though significantly inhibited the growth of velvetleaf ( Abutilon theophrasti ), did not reduce corn growth even in the presence of supplemental glycine. Wani et al . (2007) found that most of the rhizosphere isolates produced HCN in vitro and stimulated the plant growth. On the other hand, Pseudomonas entomophila showed biocontrol properties and pathogenicity due to pro­duction of HCN ( Ryall et al ., 2009 ). The Pseudomonas fragi strain CS11RH1 (MTCC 8984), produced HCN and the seed bacterization with this strain significantly increased the percentage and rate of germination, plant biomass and nutrient uptake of wheat seedlings ( Selvakumar et al ., 2009 ).

Agbodjato et al . (2015) identified five rhizobacterial species of Bacillus ( B. polymyxa, B. pantothenticus, B. anthracis, B. thuringiensis and B. circulans ), three Pseudomonas species ( P. cichorii, P. putida and P. syringae ) and Serratia marcescens . Inoculation of these rhizobacteria as biological fertilizers resulted into increased maize production. Nandi et al . (2017) found that Pseudomonas chlororaphis strain PA23 produced HCN and secreted the antibiotics pyrrolnitrin and phenazine, together with degradative enzymes and siderophores. This strain acted as a biocontrol agent. Similarly, Pseudomonas aeruginosa (HM195190) strain KC1 isolated from the rhizosphere of castor plants ( Ricinus communis ) ( Lakshmi et al ., 2015 ) was found to produce cyanide (4.78 nmol L ⁻¹ ) and seed bacterization with strain KC1 exhibited significant reduction in root length and shoot length of weed seedlings ( Amaranthus spinosus and Portulaca oleracea ) in both laboratory and glasshouse experiments. However, inoculation of strain KC1 was found less inhibitory to the seedlings of Triticum aestivum as compared to weed seedlings.

Plant pathogens produce a variety of phytotoxins that interfere with plant metabolism, ranging from subtle effects on gene expression to plant mortality ( Walton, 1996 ). Several bacterial and fungal microorganisms were also found to produce a wide array of phytotoxins with the potential to be used as herbicides ( Duke et al ., 1991 ). Two phytotoxic metabolites (prehelminthosporal and dihydropore), were isolated from the cultural filtrates of the fungus Bipolaris sp. which showed herbicidal activity against Sorghum halepense (L.) Pers. ( Parmar & Devkumar, 1993 ). The AAL-toxin (hydroxylated long-chain alkylamine containing a tricarboxylic acid moiety) produced by Alternaria alternata f. sp. lycopersici has been found to act as an effective herbicide on a range of crop and weed species. In susceptible varieties of tomatoes, it caused rapid wilting and necrosis ( Abbas et al ., 1995 ). Similarly, a phytotoxic metabolite trans-4-aminoproline isolated from culture filtrates of Ascochyta caulina was found highly effective in controlling Chenopodium album (L.) weed ( Evidente et al ., 2000 ).

Evidente et al . (2005) isolated a new phytotoxic trisubstituted naphthofuroazepinone from the culture filtrates of Drechslera siccans , named drazepinone and characterized as a 3,5,12 a trimethyl- 2,5,5a,12a-tetrahydro-1 H naphtha [2′,3′:4,5] furo [2,3- b ] azepin-2-one. The novel metabolite showed broad-spectrum herbicidal properties at 2 µg µL ^-1 solution. Another mobile phytotoxin mevalocidin, produced by Fusarium DA056446 and Roselliana DA092917 was reported to act as a broad spectrum post-emergence herbicide against grasses and broad-leaved plants ( Gerwik et al ., 2013 ). The cyclic tetrapeptide phytotoxin tentoxin produced by Alternaria alternata caused phytotoxic damage to both monocot and dicot weeds species and therefore showed the potential to be used as bioherbicide ( Saxena, 2014 ). Rath et al . (2018) investigated the role of volatile organic compounds (VOCs) produced by PGPR strains in plant growth promotion. Bacillus subtilis and Bacillus amyloliquefaciens strains produced VOCs like 3-hydroxy-2-butanone (acetoin) and 2,3-butanediol, which promoted plant growth, whereas other volatiles such as HCN and 3-phenylpropionic acid were found phytotoxic and inhibited the plant growth.

Adetunji et al . (2018) isolated an active me­tabolite mellein (a dihydroisocoumarin) from the broth of phytopathogenic strain Lasiodiplodia pseudothe­obromae and its structural characterization revealed the compound as (R)-8-hydroxy-3-methy­lisochroman-1-one. The isolated phytotoxic metabolite from Lasio­diplodia pseudotheobromae (at 10 µg µL ^-1 conc.) showed selective inhibition at 56-66% against the Poaceae and Valerianaceae families. Another bioactive phytotoxin with good herbicidal activity was extracted from Pseu­domonas aeruginosa strain C1501 and the active compound was identified as a 2-(hydroxymethyl) phenol ( Adetunji et al ., 2019 ). The C1501 strain showed significant decrease in the dry weight of Amaranthus hybridus (pig weed) seedlings. Lawrancea et al . (2019) isolated a Pseudomonas aeruginosa strain H6 with weedicide efficacy from the rhizosphere of Momordica charantia . Metabolite identified from strain H6 showed the presence of antifungal and herbicidal compounds. GC-MS analysis of the distinctive herbicidal metabolites produced by Pseudomonas aeruginosa H6 was iden­tified as quinoline derivatives, which were found highly toxic to the target weeds. Both, supernatant culture and crude extract of strain H6 showed high inhibition activity in Pennisetum purpureum , Oryza sativa , Pisum sativa and Amaranthus spinosum .

The primary mechanism of biocontrol by rhiz­obacteria involves production of antibiotics such as 2,4-diacetylphloroglucinol (DAPG), pyoluteorin, pyr­rol­­nitrin, phenazine-1-carboxyclic acid, 2-hydroxy­phe­naz­i­­nes and phenazine-1-carboxamide. Antibiotics have also been found to act as determinants in triggering in­duced systemic resistance (ISR) in the plant system and contribute to disease suppression by conferring a competitive advantage to biocontrol agents. Kataryan & Torgashova (1976) reported that the antibiotic 2,4-DAPG showed phytotoxic activity resembling to the 2,4-D herbicide. Geldanamycin and nigericin, two phytotoxic metabolites, were obtained from a strain of Streptomyces hygroscopicus and showed significant pre-emergence activity on proso millet, barnyard grass, garden cress and giant foxtail. A polyketide secondary metabolite, herboxidiene, produced by Streptomyces chromofuscus , showed potent and selective herbicidal activity against weeds but not against wheat ( Miller-Wideman et al ., 1992 ). Secondary metabolites iso­lated from Pseudomonas syringae strain 3366 were found inhibitory to downy brome and these me­tabolites consisted of phenazine-1-carboxylic acid, 2-aminophenoxazone and 2-aminophenol ( Gealy et al ., 1996 ). Similarly, phenazine-type antibiotics produced by Pseudomonas fluorescens were also reported to inhibit the root growth of downy brome weed ( Gealy et al ., 1996 ).

Bacterial species inhabiting the crop rhizosphere have been reported to affect plant growth in either a positive or in a negative way. Beneficial effects of rhizosphere bacteria have most often been based on suppression of diseases, increased seedling emergence and stimulation of plant growth along with inhibition of weeds growth (Fig. 2) ( Sindhu et al ., 2014 , 2016 ; Phour & Sindhu, 2019 ). A large array of bacteria, including species of Azospirillum , Azotobacter , Art­hrobacter , Bacillus , Enterobacter , Burkholderia , Pa­enibacillus , Pseudo­monas and Rhizobium , have been reported to enhance plant growth ( Wani et al ., 2008 ; Khan et al ., 2009 ; Sindhu et al ., 2018 ). Five bacterial isolates belonging to Pseudomonas putida (TSAU1), Pseudomonas ex­tremorientalis (TSAU6 and TSAU20), Pseudomonas chlo­roraphis (TSAU13) and Pseudomonas aurantiaca (TSAU22) were selected from the rhizosphere of wheat grown in saline soil ( Egamberdieva & Kucharova, 2009 ). These isolates produced IAA and among these four isolates caused significant increase in the shoot, root and dry matter of wheat under saline conditions. Mejri et al . (2010) reported significant gain in growth of wheat, barley, oat, pea and chickpea after inoculation with Pseudomonas trivialis strain X33d, whereas inoculation of this strain in downy brome weed caused growth inhibition.

Kennedy et al . (2001) reported stimulation of oilseed rape growth due to application of P. fluorescens strain D7, which aggressively reduced the growth of downy brome. Similarly, Li & Kremer (2006) reported increase in growth of soybean and wheat due to application of P. fluorescens strain G2-11. Previously, this strain showed suppressive effect on the growth of several weeds (barnyard grass, green foxtail and morning glory). Certain rhizosphere bacterial strains T42, L9, 7O ₀ , O ₀ 10 and W9 were found to be advantageous under field conditions by causing weed suppression and also improved the competitive ability of the crop against weeds ( Abbas et al ., 2017 ). Thus, those rhizospheric bacterial isolates that specifically colonize and inhibit growth of weeds but not that of crop plants, may be used as biological control agents. This may benefit agriculture by contributing to increased crop yields, by reducing weed competition and reducing the use of chemical herbicides ( Patil, 2014 ).

Twelve rhizobacterial isolates were tested for their effect on growth of wheat and weed under pot house conditions. Rhizobacterial isolates SYB101, CPS67 and HWM11 were found to stimulate growth of wheat and inhibited the growth of Phalaris minor ( Phour, 2012 ). Khandelwal (2016) reported that inoculation of bacterial isolate WHA87 caused 94-182% increase in RDW and 30-340% increase in SDW of wheat, whereas its inoculation showed 21-81% decrease in RDW and 33-43% decrease in SDW of Chenopodium album at 30, 60 and 90 days of plant growth under pot house conditions. In case of Asphodelus tenuifolius , inoculation of bacterial isolate MSA56 showed 231% increase in RDW and 225% increase in SDW of wheat, whereas its inoculation caused 40-85.7% decrease in RDW and 53-54.3% decrease in SDW of A. tenuifolius . Rhizobacterial isolates WHA87, MSA39, MHA75 and MSA56 were found to stimulate growth of wheat, whereas isolates MSA39 and WHA87 inhibited the growth of C. album and isolates MHA75, MHA93 and MSA56 inhibited the growth of A. tenuifolius .

In another study, rhizobacterial isolates HMM76, HMM92, JMM24, JMM35 and SYB101 were found to stimulate growth of mustard and inhibited the growth of Lathyrus aphaca under pot house conditions ( Phour, 2016 ). At 75 days after sowing, inoculation of the two bacterial isolates HMM92 and JMM24 showed 54 to 191% increase in RDW and SDW of mustard, whereas they caused 36 to 92% decrease in RDW and SDW of Lathyrus aphaca . These rhizobacterial isolates may be further tested for suppression of weed growth under field conditions for their subsequent application as bioherbicides. A better understanding of the molecular biology of plant-microbe interactions may be useful for designing of strategies in which specific microorganisms may act as PGPR for the cereal and legume crops along with suppressive effects on the growth of weeds.