Summary 449
I. INTRODUCTION 450
II. THE PARTNERS 451
1. Cyanobionts and their role 451
2. Hosts and their role 453
3. Location of cyanobionts in their hosts 455
III. INITIATION AND DEVELOPMENT OF SYMBIOSES 458
1. Initiation of symbioses 458
2. Geosiphon pyriforme 458
3. Cyanolichens 459
4. Liverworts and hornworts 460
5. Azolla 460
6. Cycads 461
7. Gunnera 461
IV. THE SYMBIOSES 462
1. Geographical distribution and ecological significance 462
2. Benefits to the partners 462
(a) Benefits to the cyanobionts 462
(b) Benefits to the hosts 463
3. Duration and stability 463
4. Mode of transmission and perpetuation 463
5. Recognition between the partners 464
6. Specificity and diversity 464
7. Symbiosis‐related genes 465
8. Modifications of the cyanobiont 466
(a) Growth and morphology 466
(b) Photosynthesis and carbon metabolism 467
(c) Glutamine synthetase 467
(d) Heterocysts 469
(e) N2fixation 470
9. Nutrient exchange 471
(a) Carbon 471
(b) Nitrogen 472
V. EVOLUTIONARY ASPECTS 472
VI. ARTIFICIAL SYMBIOSES 474
VII. FUTURE OUTLOOK AND PERSPECTIVES 475
1. Cryptic symbioses 476
2. Developmental profile of symbiotic tissues 476
3. Sensing and signalling 476
4. Genetic aspects 476
5. Physiological and biochemical aspects of nutrient exchange 477
6. Microaerobiosis 477
7. Potential applications 477
Acknowledgements 477
References 477
Cyanobacteria are an ancient, morphologically diverse group of prokaryotes with an oxygenic photosynthesis. Many cyanobacteria also possess the ability to fix N2. Although well suited to an independent existence in nature,
some cyanobacteria occur in symbiosis with a wide range of hosts (protists, animals and plants). Among plants,
such symbioses have independently evolved in phylogenetically diverse genera belonging to the algae, fungi,
bryophytes, pteridophytes, gymnosperms and angiosperms. These are N2‐fixing symbioses involving heterocystous cyanobacteria, particularly Nostoc, as cyanobionts (cyanobacterial partners). A given host species
associates with only a particular cyanobiont genus but such specificity does not extend to the strain level. The
cyanobiont is located under a microaerobic environment in a variety of host organs and tissues (bladder, thalli and
cephalodia in fungi; cavities in gametophytes of hornworts and liverworts or fronds of the Azolla sporophyte;
coralloid roots in cycads; stem glands in Gunnera). Except for fungi, the hosts form these structures ahead of the
cyanobiont infection. The symbiosis lasts for one generation except in Azolla and diatoms, in which it is
perpetuated from generation to generation. Within each generation, multiple fresh infections occur as new
symbiotic tissues and organs develop. The symbioses are stable over a wide range of environmental conditions,
and sensing–signalling between partners ensures their synchronized growth and development. The cyanobiont
population is kept constant in relation to the host biomass through controlled initiation and infection, nutrient supply and cell division. In most cases, the partners have remaine...